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BIOSTRATIGRAPHY AND DIVERSITY PATTERNS OF 
CENOZOIC ECHINODERMS FROM FLORIDA 


By 

CRAIG W. OYEN 


A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL 
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT 
OF THE REQUIREMENTS FOR THE DEGREE OF 
DOCTOR OF PHILOSOPHY 

UNIVERSITY OF FLORIDA 


2001 


I dedicate this to my parents, Walter and Norma Oyen, who have always 
expressed absolute confidence in my ability to succeed and supported all my 
endeavors without question or hesitation. I could not have done this without their 
support through all these years and I appreciate it more than they realize. 


ACKNOWLEDGMENTS 


This project could not have been completed without the aid of many 
people. Most important is the assistance and direction given by my dissertation 
committee chair, Dr. Douglas S. Jones. He encouraged me to work on whatever 
topic(s) I found interesting, and simply gave me suggestions to improve my 
approach in order to answer any of those questions. He also made my time in 
Gainesville enjoyable academically and socially by introducing me to other 
faculty and students, inviting me to his home or restaurants for dinners, and 
participating in pick-up basketball games and intramural games for relaxation. 

He (along with Roger Portell) accompanied me on many fascinating fieldtrips in 
Florida and other locations (often in association with GSA meetings) that 
expanded my scientific and other perspectives greatly. I thank the other 
members of my dissertation committee, Drs. Randazzo, Hodell, MacFadden, and 
Mature, for participating in my research and providing guidance whenever I 
asked for their help. I consider it a pleasure to have had this group of faculty 
members participating on my committee because they only treated me with 
respect and they openly provided suggestions they believed would serve me best 
in the context of completing my research and degree. 


I also thank Roger Portell (Collection Manager, IP Division, FLMNH) for 
his extensive help in most aspects of this dissertation. He spent much time with 
me during all phases of the research to help me become familiar with the 
invertebrate paleontology collection in the FLMNH, aided in nearly all fieldwork 
excursions, and accompanied me on visits to other museums or personal fossil 
collections. He also supplied me with many critical publications from his personal 
library and found others that I was able to purchase for myself. His contacts with 
non-professional fossil collectors in Florida enabled me to gather data I may not 
have been able to access otherwise. Roger’s help lessened some of the stress 
associated with my research. 

Financial assistance to complete the dissertation was provided by many 
sources. Grant proposals I submitted were funded by the Geological Society of 
America, the American Federation of Mineralogical Societies, Sigma Xi, the 
R. Jerry Britt Jr. Paleobiology Award (of the Florida Museum of Natural History), 
the Gary S. Morgan Student Research Award (of the Florida Paleontological 
Society), and the Mitchell Hope Scholarship Award (of the Southwest Florida 
Fossil Club). Employment by the Department of Geological Sciences provided 
teaching assistantships (under Dr. A. F. Randazzo, Chair) and research 
assistantships (under Dr. G.H. McClellan). The Florida Museum of Natural 
History provided research assistantships (under Dr. D.S. Jones). Members of my 
family also contributed significant financial assistance to my endeavors, 
particularly my parents (Walter and Norma Oyen) and my sister (Valerie Oyen- 
Larsen). 


IV 


I am very grateful for the thoughtfulness of Barbara and Reed Toomey, 
Roger and Anne Portell, Douglas and Sheila Jones, Jewel Pozefsky, Kendall 
Fountain, and Richard Hulbert. They gave me a place to stay (usually for 
extended periods of time) while traveling from Georgia or Pennsylvania during 
the holidays, long weekends, or summer breaks, so that I could finish my work. 

Fieldwork assistance was provided by many people during both the 
modern echinoderm study and fossil echinoderm collection phase of the 
dissertation. I thank Dr. Frank Mature (Zoology Department) for giving me 
guidance, equipment, and access to Seahorse Key during my study of the 
modern echinoids around the island. Dr. Douglas Jones, Roger Portell, and 
Kevin Schindler (FLMNH) helped construct the enclosures on site and also 
accompanied me during periodic visits to gather data. Field assistance on this 
part of the project (in addition to that listed above) was provided frequently by 
Karen Powers, Rich and Nicole Hisert, Len Fishkin, Kendall Fountain, and Ross 
Russell (all fellow students at UF). Assistance with fossil collecting was 
dominated by Dr. Douglas Jones, Roger Portell, and Kevin Schindler (of the 
FLMNH). 

A number of individuals made their personal fossil collections available for 
examination or donated specimens to the Florida Museum of Natural History that, 
in turn, allowed me to incorporate the information into my dissertation. I would 
like to thank those people who provided a significant number of specimens. The 
individuals with an academic or research affiliation have that information included 
in the parentheses after their name: Dr. Burchard Carter (Georgia Southwestern 


V 


state University), Dr. Jonathon Bryan (Okaloosa-Walton Community College), 
Dr. Thomas Scott (Florida Geological Society), Drs. Emily and Harold Vokes 
(Tulane University), Dr. Lyle Campbell (University of South Carolina at 
Spartanburg), Harley Means (Florida Geological Survey), Dr. Sherwood Wise 
(Florida State University), Muriel Hunter (formerly of Coastal Petroleum Co.), 
Jules DuBar, Phil Whisler, Tim Cassady (deceased), Byron Shumaker, Wendy 
Conway, Gary Schmelz, and Charles Hewlett. 

Access to quarries and property is not a simple process any longer, and 
several mine operators or owners have been generous in permitting me and 
other paleontologists to collect from their sites. This study benefited from the 
assistance given by Larry Rogers (Limestone Products, Inc.), C.T. Williams 
(Florida Rock Industries), Tom Jones (formerly of DoLime Products, Inc.), Fred 
Pirkle (formerly of DuPont Corporation), Richard Brown (Quality Aggregates, 
Inc.), Hugh Cannon (formerly of Quality Aggregates, Inc.), and Jimmy Philman 
(Handyphil, Inc.). 

1 appreciate assistance in completing the photographic work for the 
dissertation document, particularly when I was nearing completion of the 
manuscript and didn’t have time to do it myself. Two people, in particular, 
allowed this to occur more easily and rapidly: George Hecht (FLMNH) 
photographed many of the fossils and Terry Lott (FLMNH) printed most of the 
photos. 

I thank many of my fellow students at the University of Florida for 
providing entertainment and camaraderie while engaged in the academic, 


VI 


athletic, and other endeavors during my time here. In particular, this includes all 
the “Psychotic Basement Troll” intramural basketball team players (Jose Garrido, 
Kendall Fountain, Len Fishkin, Greg Ferris, Rich Hisert, Joe Stoner, Stan 
Crownover, Robin Graves, Chris Saum, Neil Johnson, and Dr. Douglas Jones, 
among others) over the years. We weren’t pretty, but we had fun. My physicians 
benefited financially from this activity as well, as a result of my multiple sports 
injury visits. 

Finally, I thank my parents (Walter and Norma), brothers (Lance and 
Mitch), and sister (Valerie) for never asking “why” I was doing the research, and 
only asked” when” they could come for graduation. 


VII 


TABLE OF CONTENTS 


page 

ACKNOWLEDGMENTS iii 

KEY TO SYMBOLS xi 

ABSTRACT xiii 

CHAPTERS 

1 INTRODUCTION 1 

General Overview 1 

Previous Work 2 

Purpose and Goals 5 

Methods and Materials Studied 8 

2 STRATIGRAPHY OVERVIEW AND GEOLOGIC SETTING 11 

Introduction 'll 

Eocene Stratigraphy 12 

Oligocene Stratigraphy 14 

Miocene Stratigraphy 14 

Pliocene Stratigraphy 16 

Pleistocene Stratigraphy 17 

3 SYSTEMATIC PALEONTOLOGY OF FLORIDA ECHINODERMS 19 

Introduction 19 

Class Echinoidea Fossils 21 

Eocene Echinoids 21 

Oligocene Echinoids 92 

Miocene Echinoids 108 

Pliocene Echinoids 150 

Pleistocene Echinoids 220 


viii 


Class Crinoidea Fossils 237 

Lower Ocala Limestone Crinoids 237 

Upper Ocala Limestone Crinoids 239 

Class Asteroidea Fossils 241 

Eocene Asteroids 241 

Oligocene Asteroids 242 

Miocene Asteroids 243 

Pliocene Asteroids 244 

Class Ophiuroidea Fossils 247 

Eocene Ophiuroids 247 

Miocene Ophiuroids 248 

Pliocene Ophiuroids 249 

4 ECHINODERM DIVERSITY PATTERNS AND BIASES 259 

Taxonomic and Biostratigraphic Discussion 259 

Eocene Echinoids 260 

Oligocene Echinoids 265 

Miocene Echinoids 268 

Pliocene Echinoids 272 

Pleistocene Echinoids 276 

Crinoids 277 

Asteroids 279 

Ophiuroids 282 

Biases in the Cenozoic Echinoderm Record 284 

Resolution of Data 284 

Stratigraphic Nomenclature 285 

Outcrop Exposure and Relief 286 

Carbonate Versus Siliciclastic Environments 288 

Age of Stratigraphic Units 292 

Epoch Duration 292 

Taxonomic Nomenclature 295 

Collector Bias 298 

Substrate and Facies Preferences of Echinoids 299 

The Eocene-Oligocene Diversity Change 308 

Early Paleogene Oceanographic Conditions 310 

Early to Middle Paleogene Transitions 311 

Late Eocene Extinctions and Biogeographic Patterns 314 

Chapter Summary 322 

5 ALLOMETRIC HETEROCHRONY IN MELLITID ECHINOIDS: A CASE 

STUDY FROM FLORIDA 325 

Preface to the Biometric Analysis 325 

Original Research Objectives 325 

Growth Study Procedure 326 


IX 


Introduction and Heterochrony Overview 329 

Materials and Methods 333 

Materials Examined 333 

Data Acquisition Methods 334 

Biometric Traits Evaluated 336 

Data Analysis Methods 339 

Results 340 

6 SUMMARY AND CONCLUSIONS 349 

New Echinoderm Diversity Patterns 349 

Taxonomic Implications 352 

What Work Lies Ahead? 354 

APPENDIX 356 

REFERENCES 421 

BIOGRAPHICAL SKETCH 436 


X 


KEY TO ABBREVIATIONS 


FLMNH 

UF 

UF# 

USNM 

USNM# 

USGS 

FGS 

TL 

TW 

PSL 

PSW 

PSP 

PPL 

PPW 

PPP 

POSAP 

ANLL 

ANEW 

ANLP 

PD(I-V) 

TWMX 

LTH(1-5) 

TTH(1-5) 

PAL(I-V) 


PAW(I-V) 


THMX 


Florida Museum of Natural Flistory. 

University of Florida. 

Invertebrate Paleontology fossil lot number, Florida Museum of 
Natural History, University of Florida. 

United States National Museum. 

United States National Museum paleontology fossil lot number. 
United States Geological Survey. 

Florida Geological Survey. 

Test length (along anterior to posterior transect). 

Test width (at midpoint of TL). 

Peristome length (along anterior to posterior transect). 

Peristome width. 

Peristome position (from anterior test margin to anterior peristome 
margin). 

Periproct length. 

Periproct width (at midpoint of PPL). 

Periproct position (from anterior periproct margin to anterior 
peristome margin). 

Position of apical system (from anterior test margin to center of 
apical system). 

Anal lunule length (interior lunule length, along anterior to posterior 
transect on aboral side). 

Anal lunule width (interior lunule width on aboral side, at midpoint of 
lunule length). 

Anal lunule position (distance from anterior lunule margin to apical 
system center). 

Pressure drainage channel span (maximum width, adoral surface; 
Roman numerals l-V indicate Loven ambulacral designations). 

Test width maximum. 

Longitudinal test height (at five equidistant points beginning at 
anterior test margin and proceeding toward posterior). 

Transverse test height (at five equidistant points starting at left test 
margin when viewing adapical surface from above). 

Petaloid ambulacrum length (aboral ambulacrum length, from apical 
system margin to maximum pore-pair position; Roman numerals 
indicate Loven designations for ambulacra). 

Petaloid ambulacrum width (aboral ambulacrum width, at midpoint 
of ambulacrum length, from outer pore-pair to outer pore-pair; 
Roman numerals indicate Loven designations for ambulacra). 

Test height maximum. 


XI 


AL(I-V) 

IL(1-5) 

ALL(I-V) 

ALW(I-V) 

ALP(I-V) 


Ambulacrum length (adoral surface ambulacrum, from peristome 
margin to test margin; Roman numerals indicate Loven 
designations for ambulacra). 

Interambulacrum length (adoral surface interambulacrum, from 
peristome margin to test margin; Arabic numerals indicate Loven 
designations for interambulacra). 

Ambulacral lunule length (measured on aboral surface using interior 
length; Roman numerals indicate Loven designations for 
ambulacra). 

Ambulacral lunule v\/idth (measured on aboral surface, interior width 
at midpoint of length; Roman numerals indicate Loven designations 
for ambulacra). 

Ambulacrum lunule position (measured from center of apical system 
to adapical lunule margin; Roman numerals indicate Loven 
designations for ambulacra). 


XII 


Abstract of Dissertation Presented to the Graduate School 
of the University of Florida in Partial Fulfillment of the 
Requirements for the Degree of Doctor of Philosophy 

BIOSTRATIGRAPHY AND DIVERSITY PATTERNS OF 
CENOZOIC ECHINODERMS FROM FLORIDA 

By 

Craig W. Oyen 
May 2001 

Chair: Douglas S. Jones 

Major Department; Geological Sciences 

Fossil echinoderms from the Middle Eocene through Pleistocene in Florida 
are described here in detail, including any previously reported species as well as 
taxa that are new records as a result of this study. Twenty-three formations from 
the state produced fossils belonging to four classes of echinoderms, including 
echinoids, asteroids, crinoids, and ophiuroids. Echinoids are distinctly more 
diverse and documented than the other classes in the state. 

The diversity of echinoderms from the Cenozoic in Florida has increased 
significantly as a result of examination of fragmented skeletal debris and small 
size fractions of the sedimentary rock. Echinoid species diversity generated 
herein resulted in an increase from 68 species (reported in publications before 
1994) to a current diversity of 97 species. These new additions include both 
newly reported stratigraphic records of a species from the state and new 
taxonomic records (i.e., undescribed species). An important result of the 
additional taxa recorded is the change in the echinoid diversity pattern over time. 


Most newly reported taxa were found in Miocene (14 new records) and Pliocene 
(eight new records) formations. The diversity pattern thereby shows a dramatic 
increase in diversity from the Oligocene to the Miocene, rather than a drop in 
diversity as formerly documented. 

Changes in the diversity pattern may be attributed to multiple biases in 
addition to species extinction and origination. Such biases include stratigraphic 
resolution of data, changes in stratigraphic nomenclature, amount of outcrop 
exposure and topographic relief, varying preservation potential in carbonate 
versus siliciciastic strata, unequal epoch duration in the Cenozoic units, 
taxonomic nomenclature changes (i.e., species versus subspecies 
identifications), fossil collector preferences favoring unbroken specimens, and 
substrate grain-size preferences of the echinoderm taxa. Analysis of such biases 
indicated collector bias, strata composition, and substrate texture were the most 
important factors ultimately controlling the diversity pattern in Florida’s Cenozoic 
echinoderm record. 

Finally, an analysis of allometric heterochrony patterns completed for 
echinoid species in the family Mellitidae showed the predominance of 
paedomorphic patterns of evolution. Such trends closely matched previously 
proposed models for r-selection styles in higher energy environments where 
descendant species ultimately inhabited. 


XIV 


CHAPTER 1 
INTRODUCTION 

General Overview 

The fossil record of Florida is diverse and rich, particularly with regard to 
marine invertebrate groups such as the mollusks, echinoderms, and foraminifera; 
and the terrestrial mammals. Echinoderms in particular are usually well 
preserved, abundant, and an important component of Florida’s Cenozoic fossil 
record. These fossils have been studied and documented in varying degrees for 
approximately 150 years, though no comprehensive study of all fossil 
echinoderms in the state has been completed until now. It is my objective in this 
study to present a taxonomic overview of all echinoderms from the Middle 
Eocene through the Pleistocene, and to provide specific diversity values (at 
stratigraphic unit and epoch level resolution) for this time range. Many taxa are 
discussed at the generic level herein because most of my new records are not 
yet described to species level and, furthermore, numerous taxa have subspecies 
that I believe need to be re-evaluated. New species descriptions will be 
published as part of a continuing program of investigation. 


1 


2 


Previous Work 

The echinoderms of Florida have been documented as parts of 
monographs or descriptive papers from various Cenozoic stratigraphic units. 

Most of the early papers were limited in their breadth of discussion to the 
echinoids however, with no references to other groups of echinoderms that were 
part of the formations or, at best, only general statements to the effect that sea 
star ossicles or possible crinoid ossicles were present. One of the first 
monographs to include a detailed description of known Florida taxa was by Clark 
and Twitchell (1915), in which they discussed all records of Mesozoic and 
Cenozoic echinoderms from the U.S.A. At the time of this work, relatively few 
echinoderms had been reported formally or described from Florida, so this 
monograph contained few records for the state. 

Echinoids from Florida and other southeastern states were the focus of 
several of Cooke’s major monographic works (e.g., 1941, 1942, 1959), including 
the most comprehensive biostratigraphic work on echinoids in the southeastern 
U.S.A. to date (Cooke, 1959). A geologist with the U.S. Geological Survey, 
Cooke worked extensively in the Coastal Plain of the southeastern U.S. 
describing the sedimentary rock, strata, and fossils in this region (including 
Florida). His work helped generate a foundation for both the paleontologic and 
stratigraphic framework of the state (e.g., Cooke, 1915; Cooke and Mossom, 
1929; Cooke and Mansfield, 1936; Cooke, 1939; Cooke, 1945). Durham focused 
primarily on the western U.S. fossils and localities, but two of his papers 
(Durham, 1954; 1955) included significant references to Florida echinoids as part 


3 


of his revision and updates of the order Clypeasteroida. Fischer (1951) 
described the echinoid fauna found in the Lov\/er Ocala Limestone (the 
stratigraphic unit formerly referred to as the Inglis Formation). Echinoderm 
microfossils, including the comatulid crinoids, were discussed by Howe (1942) as 
being a neglected group of fossils in the Gulf Coastal Plain region, but he did not 
figure or refer to any Florida crinoids in his work. 

During the middle to late twentieth century, a new group of echinoderm 
paleontologists continued to collect and describe fossils from Florida and the 
surrounding states. One of the most prolific of these workers was Porter Kier. 
Kier published numerous papers and monographs on stratigraphic occurrences 
and taxonomic descriptions of echinoids from Florida, as well as other areas of 
the southeastern Coastal Plain and the Caribbean that have relevance to the 
Florida taxa. Among his numerous monographs, several have specific relevance 
to Florida fossils and particular value in taxonomic and biostratigraphic work, 
including 1) a revision of the cassiduloid echinoids (Kier, 1962); 2) descriptions of 
the Caloosahatchee Formation and Tamiami Formation echinoids of Florida 
(Kier, 1963); 3) a revision of the oligopygoid echinoids (Kier, 1967); 4) a 
description of Middle Eocene echinoids of Georgia, that covers concurrent 
species occurrences in Florida (Kier, 1968); 5) a descriptive work on the 
spatangoid echinoids of Cuba that, once again, involves species that are present 
in Florida as well (Kier, 1984); and finally 6) a discussion of life habits of modern 
taxa in the Florida Keys, that includes observations relevant to fossil taxa in the 
state (Kier and Grant, 1965). 


4 


More recently, fossil echinoderm work involving non-echinoid taxa in 
Florida included analyses of new occurrences of asteroids from the Eocene and 
Pliocene (Jones and Portell, 1988; Ivany et al., 1990); new records of juvenile 
ophiuroids found as fossils within seagrass concentrations in Eocene limestones 
(Ivany et al., 1990); and the first confirmed record of comatulid crinoids for the 
state (Oyen, 1995). Studies of echinoid substrate preferences and their 
associated distributions in Florida were published by Carter (1987, 1990, 1997), 
Carter et al. (1989), Carter and McKinney (1992), and McKinney and Zachos 
(1986), while new taxonomic records and stratigraphic occurrences were 
published by Oyen and Portell (1996) and Portell and Oyen (1997). Finally, the 
use of echinoids for biostratigraphic purposes has been discussed within the 
context of Eocene stratigraphy. Flunter (1976) suggested that based on her 
extensive fieldwork, she believed that the oligopygoid echinoids might serve as 
viable biostratigraphic markers for the Late Eocene limestones in Florida. Shortly 
thereafter, Zachos and Shaak (1978) formally proposed a biozonation based on 
the stratigraphic ranges of the three Florida species of Oliqopygus , which 
resulted in becoming the best known and most widely used echinoid biozone for 
the state’s strata. 

Unfortunately, no attempt was made thus far to assimilate all the 
information regarding Florida records into a single paper. Therefore, it is my goal 
in the following sections to familiarize paleontologists with the current status of 
diversity values for all strata exposed at the surface in the state. I also interpret 


5 

diversity patterns and provide information regarding possible biases or sampling 
artifacts for the echinoderm diversity data in Florida. 

Purpose and Goals 

The primary focus of this dissertation is to improve the fundamental 
knowledge regarding stratigraphic distribution of all varieties of fossil 
echinoderms of Florida. As noted earlier, significant work describes and 
documents fossil echinoids in Cenozoic rocks of the state, but such work has 
slowed dramatically in recent decades. This means that the database for 
interpretations of diversity, extinction and speciation patterns, and paleoecology 
has become stagnant. Although topics of greater interest in the discipline of 
paleontology have shifted in recent years away from classic paleontology studies 
such as taxonomy and biostratigraphy, I believe it is important to continue to 
expand the essential knowledge of species diversity. Such data are critical to 
build models and solutions explaining any trends over time. The best 
interpretations and predictions can only be generated if work continues to build 
on the alpha-level taxonomy and diversity database; the “big picture” is 
absolutely dependent on the smaller pieces of the puzzle to make the picture 
clear. While collecting echinoderms over the past fifteen years, it has become 
obvious to me that many new species of echinoids, asteroids, crinoids, and 
ophiuroids are part of the fossil record in Florida. Although I am not proposing 
new species names for any “new” taxa that I or others have collected, I am 
providing detailed descriptions of these fossils to be used as the basis for 


6 


comparison and in preparation for the formal review process of publication after 
this dissertation. Therefore, my research is directed at providing the most current 
and most thorough description and discussion of taxa in the state since the work 
of Cooke (1959) and the works of Kier (mostly in the 1 960s). 

A second goal of my research is to interpret the diversity pattern for the 
echinoderms of Florida, with particular focus on the echinoids. How has the 
diversity pattern changed in comparison with what was previously known and 
published? What implications do any of these changes have for understanding 
the basis for trends in the diversity over time? All paleontologists recognize that 
taphonomic processes greatly affect their ability to clearly read the fossil record 
and interpret information such as ecological conditions or physiological 
characteristics of those former organisms of interest. Therefore, I provide 
possible reasons why the diversity pattern looks as it does by accounting for 
specific biases affecting the fossils and strata of Florida. As an example, the 
Paleogene stratigraphic units are predominantly carbonate rocks whereas the 
Neogene units are dominantly siliciclastic rocks. Preservation of fossils within 
the carbonate units tends to be better than that observed in the siliciclastic units. 
The diagenetic changes occurring in carbonates may be less destructive toward 
fossils in contrast to when they are contained within siliciclastic units. Does this 
situation have a significant impact on the diversity record of echinoderms in 
Florida, and if so, why? Questions such as this are explored and addressed in 
my research presented herein to fully explain the patterns that exist in the data. 


7 


Another goal of this dissertation is to provide a thorough and current 
summary of the nomenclature history, lithology, and physical characteristics of all 
of the stratigraphic units that contain the fossil echinoderms of interest. It is not 
my intent to justify or solve the myriad problems and disputes associated with 
various formations in the state. However, it is valuable to include these 
descriptions, at least from a practical perspective, so that the information is 
available for interested workers who may refer to this dissertation in the future. I 
prefer to consider this dissertation work to be a comprehensive monograph on 
the echinoderm paleontology and stratigraphy for the state. Inclusion of details 
regarding the sedimentology and stratigraphy allows my research to be a 
functional reference source for other researchers interested in Florida geology 
and paleontology. 

Finally, a component of my research was directed at interpreting the 
allometric heterochrony patterns that exist in closely related Neogene and 
Recent species of echinoids of the family Mellitidae. McKinney produced most of 
the evolutionary interpretations for echinoids of Florida and the Caribbean to date 
(e.g., McKinney, 1984 and 1986, among others), though most of his analyses 
focused on Paleogene taxa. Are the patterns present in the Paleogene echinoids 
also present in the mellitids (that evolved in the Neogene)? Are the heterochrony 
patterns resulting from ecology-driven change similar to those patterns indicated 
by the Paleogene taxa? These were among the questions I considered when 
analyzing the fossil mellitid species of the Pliocene through the Recent strata of 
the state. The analysis of biometric data from the included species also provided 


8 


further insight into the taxonomic status of selected species of echinoids, such as 
Mellita guinquiesperforata , and the quantitative validity of proposed qualitative 
characteristics of the taxa. 

Methods and Materials Studied 

Fossils included in the database for this study were obtained from a 
variety of sources. I conducted fieldwork and collected Florida echinoderms for 
nearly 15 years, and incorporated the information gathered before and during my 
graduate studies at the University of Florida into this dissertation. As part of this 
research, more than 100 echinoid-bearing localities were sampled, including 
many sites that are no longer accessible. All specimens found during my study 
are now part of the research collection in the Invertebrate Paleontology (IP) 
Division, Florida Museum of Natural History (FLMNH), University of Florida 
(collection acronym UF), Gainesville. The IP Division at the FLMNH holds the 
largest number of fossil echinoderms from Florida. This collection serves as the 
primary data source for my work, as well as for other Cenozoic echinoderm 
workers in the southeastern U.S.A. 

Additional sources of data regarding Florida fossil echinoderms include 
other museum and private collections and published records. I examined 
specimens in the Department of Paleobiology, U.S. National Museum (USNM) at 
the Smithsonian Institution; the Florida Geological Survey; materials collected 
and curated by Harold and Emily Vokes, formerly of the Department of Geology, 
Tulane University; the Jules DuBar collection; the collection of Burt Carter at 


9 


Georgia Southwestern State University; the Florida State University Geology 
Department collection; and the stratigraphic materials collected by Joe Banks 
and Muriel Hunter (formerly of the Coastal Petroleum Company) and the 
collections were used to assimilate my echinoderm data. Each of these listed 
collections (with the exception of the B. Carter and USNM collections) are now 
part of the IP Division of the FLMNH. Finally, many individual collectors donated 
specimens to the FLMNH and greatly improved my ability to add new information 
to the Florida fossil echinoderm database. 

Some of the taxa collected were found by sieving micro-size fractions of 
poorly lithified sedimentary rocks, as well as through the production of silicone 
peels of external molds within the sedimentary rocks. Relatively few stratigraphic 
sections or localities were examined closely at the small sediment or fossil size 
range, and this provided some of the new records reported in this paper. To 
date, only two stratigraphic units (or portions of these units) were so sampled: the 
Lower Ocala Limestone and the Chipola Formation. Work continues in the 
FLMNH IP Division to sieve additional strata as well as to produce many more 
silicone peels to augment the original rock samples and fossils collected in the 
field. 

Details regarding the methods applied to the biometric analyses for 
heterochrony interpretations are included in Chapter 5. However, the specimens 
representing the variety of mellitid species considered in that chapter were 
obtained from fieldwork, museum collections including those of the FLMNH IP 
Division and the Invertebrate Paleontology division of the USNM. All of the 


10 


specimens of modern echinoids examined and measured were collected during 
my fieldwork at the University of Florida’s marine biology field station at Seahorse 
Key, located approximately 5 km southwest of Cedar Key, Florida, in the Gulf of 
Mexico. Additional background information and specific methods of data 
gathering are included where appropriate within subsequent chapters. 


CHAPTER 2 

STRATIGRAPHY OVERVIEW AND GEOLOGIC SETTING 

Introduction 

The stratigraphy of Florida is complex, partly due to the history of the 
evolving stratigraphic nomenclature and its relationship to fossil components in 
the rock. The purpose of this section is not to re-define Florida stratigraphy, but 
rather to synthesize the data regarding echinoderm-bearing formations and their 
compositions. The objective of this approach is to enhance our understanding of 
fossil echinoderm distribution in time and space. Therefore, I provide a general 
statement about each of the formations in Florida where echinoderms are found, 
along with references where interested readers may find additional information. 
Knowledge of the stratigraphic nomenclature used in the state plays a critical role 
in interpreting the biostratigraphic data presented herein, and thus is essential 
background information for this dissertation. The number of new stratigraphic 
and taxonomic records for the formations of Florida depends on how stratigraphic 
nomenclature is interpreted in the state. Unfortunately, many of the formations in 
the state were defined (at least in part) based on fossil content. This created 
problems according to the current guidelines for naming strata, i.e., the North 
American Stratigraphic Code (NACSN, 1983). Therefore, this chapter includes 
general statements regarding the lithology of formations and multiple references 


11 


12 

for literature relevant to each of the formations included in my research. This 
provides at least a basic context for the biostratigraphy discussion that follov\/s. 

Eocene Stratigraphy 

Echinoderms are present in both of the Eocene formations that are 
exposed at the surface at some location in the state (Figure 2-1). The Eocene 
strata have particular significance for analysis of echinoderm diversity because 
this highest diversity is recorded from these formations. The Middle Eocene 
Avon Park Formation is the oldest rock unit exposed at the surface in Florida. A 
second echinoderm-bearing formation, the Late Eocene Ocala Limestone, 
unconformably overlies the Avon Park. These carbonate units produce the most 
diverse echinoderm fauna of any stratigraphic interval in Florida, with the Ocala 
Limestone containing nearly all of the recorded taxa. 

The formations are very pure carbonates (especially the Ocala Limestone) 
ranging from wackestones to grainstones, and non-carbonate grains rarely 
exceed 5-10% of the rock volume. The combined thickness of the formations 
ranges from only a few m to over 365 m in the subsurface (Chen, 1965). The 
stratigraphic nomenclature associated with these Eocene formations is still under 
debate, but the names provided herein are currently accepted as valid by the 
Florida Geological Survey and the U.S. Geological Survey. For more detailed 
descriptions of these units, see Dali and Harris (1892), Cooke (1915), Applin and 
Applin (1944), Vernon (1951), Puri (1953a), Puri (1957), Chen (1965), Hunter 
(1976), Jones (1982), Scott (1991), Oyen (1995), and Randazzo (1997). 


13 


EPOCH 

STRATIGRAPHIC UNITS 

CO 

Satilla Formation 

LU 

Q. 

Anastasia Formation 

Bermont Formation 

PLIO. 

1 

Nashua 

Formation 

Caloosahatchee 

Formation 

Intracoastal 

Formation 

Jackson Bluff Tamiami 

Formation Formation 

LU 

Z 

LU 

o 

o 

Peace River Formation 

Shoal River 
Formation 

Statenville 

Formation 

Coosawhatchie 

Formation 

Torreya 

Formation 

Chipola 

Formation 

Marks Head 
Formation 

Chattahoochee 

Formation 

Parachucia 

Formation 

Arcadia 

Formation 

ono 

Bridgeboro Limestone 

Marianna Limestone 

Suwannee Limestone 

LU 

Z 

LU 

O 

O 

LU 

Ocala Limestone 

Upper Member 

Lower Member 

Avon Park Formation 


Figure 2-1 . Middle Eocene through Pleistocene stratigraphic units containing 
echinoderm fossils. This illustration is schematic only and is not 
intended to relate information regarding unconformities, hiatuses, or 
facies. Formations without echinoderms are not included here. 


14 


Oligocene Stratigraphy 

The echinoderm-bearing Oligocene rocks of Florida include the Suwannee 
Limestone, the Marianna Limestone, and the Bridgeboro Limestone (Figure 2-1). 
These three formations are similar to the Eocene units in their carbonate-rich 
composition, although the non-carbonate mineral content may exceed 10% 
slightly more frequently than is true for the older strata. Lithologies of the strata 
range across the spectrum from mudstones to grainstones (Bryan, 1991), and 
also vary from poorly lithified facies to very well-cemented or partially silicified. 
The most pervasive stratigraphic unit is the Suwannee Limestone, and all 
echinoderms from the Oligocene are present in this formation. The thickest unit 
is the Suwannee Limestone, reaching a maximum of approximately 46 m in 
northern Florida and southern Georgia along the Gulf Trough (Bryan, 1991). 

For additional information on stratigraphic nomenclature history, lithology, 
and distribution of these formations, readers should refer to Dali and Harris 
(1892), Guppy and Dali (1896), Matson and Clapp (1909), Cooke (1915), Cooke 
and Mossom (1929), Bryan (1991), Bryan and Huddlestun (1991), Huddlestun 
(1993), and Randazzo (1997). 

Miocene Stratigraphy 

Most of the research completed in recent years regarding Miocene 
lithostratigraphy for the states of Florida and Georgia was published by Thomas 
Scott (Florida Geological Survey) and Paul Huddlestun (formerly of the Georgia 
Geological Survey). Both Scott and Huddlestun produced detailed bulletins for 


15 


their respective state geological surveys in 1988, reviewing the history and 
current status of Miocene stratigraphy in Florida and Georgia. Even though their 
work improved and clarified the use of stratigraphic terminology within the two 
states, some points of debate still continue regarding the revisions they 
proposed. However, these disputes or scientific problems with stratigraphic unit 
names or lithologic definitions are not addressed in this dissertation. 

Ten Miocene formations contain echinoderms within the state of Florida 
(Figure 2-1). Formation definitions and boundaries I use in this dissertation 
currently are accepted as valid units by the Florida Geological Survey. The large 
number of stratigraphic units prevents a detailed discussion of their composition 
and areal distribution, but references cited below contain such information. In 
general, the dominant lithology of the Miocene formations is more strongly 
siliciclastic in contrast to the older Oligocene and Eocene formations. Several of 
the Miocene units contain abundant carbonate beds, while others contain few 
carbonate-rich zones and primarily consist of grains of quartz, chert, and various 
clay and phosphate minerals. The total thickness of Miocene sediments exceeds 
100 m in local areas of the subsurface of Florida (Scott, 1997). 

Specific details regarding lithology, fossil content, areal distribution, and 
the historical evolution of stratigraphic nomenclature for the Miocene formations 
are available in previously published work. In addition to Huddlestun (1988) and 
Scott (1988), these works (listed chronologically) include; Langdon (1889), 
Foerste (1893), Dali and Harris (1892), Dali and Stanley-Brown (1894), Sloan 
(1908), Matson and Clapp (1909), Veatch and Stephenson (1911), Gardner 


16 


(1926), Cooke and Mossom (1929), Cooke (1936), Mansfield (1937), Cooke 
(1943), Cooke (1945), Parker (1951), Puri (1953b), Puri and Vernon (1964), 
Heron et al. (1965), Brooks (1966), Hunter (1968), Banks and Hunter (1973), 
Abbott (1974), Wilson (1977), King and Wright (1979), Huddlestun and Hunter 
(1982), Jones and Portell (1988), Portell (1989), Yokes (1989), and Scott (1997). 

Pliocene Stratigraphy 

The Pliocene stratigraphy of Florida is extremely complex and still far from 
clearly defined and accepted by all researchers working on these units. The 
unifying theme throughout the Pliocene, just as it is through many of Florida’s 
Cenozoic intervals, is the use of fossils to help identify the formations. It is 
inevitable that formation descriptions include paleontological discussions, 
because some of the world’s richest and most densely packed fossil beds are 
found in Florida strata (e.g., the Caloosahatchee Formation of south Florida). In 
some cases, bioclasts are more than 75% of the sediment component of the 
beds and biostratigraphers used fossil species to aid in stratigraphic descriptions. 
Therefore, stratigraphic boundaries and descriptions were debated in the past, 
and likely will continue in the future as research proceeds on the stratigraphy of 
the Pliocene Epoch in the state. This time interval has many fossil-rich zones, 
and five Pliocene formations contain fossil echinoderms (Figure 2-1). 

The Pliocene units range in composition from dominantly quartz sand 
beds, to carbonate-rich layers, to shell beds with little matrix. Variation in 
lithofacies occurs within the formations, but in general, these units tend to be 


17 


higher in siliciclastic content than the Paleogene formations. Thickness of the 
Pliocene units also varies significantly, ranging from only a few meters (in surface 
and subsurface intervals) to over 100 m in the thickest sections (subsurface 
intervals only). Additional information regarding stratigraphic nomenclature 
history, lithology and unit descriptions, and fossil content is available from 
multiple sources. Examples of these (in chronological sequence) include: 

Heilprin (1887), Langdon (1889), Dali and Harris (1892), Matson and Clapp 
(1909), Sanford (1909), Mansfield (1924), Cooke and Mossom (1929), Mansfield 
and Ponton (1932), Mansfield (1939), Parker and Cooke (1944), Cooke (1945), 
Parker (1951), Puri (1953b), Dubar (1958), Dubar and Taylor (1962), Puri and 
Vernon (1964), Hunter (1968), Dubar (1974), Huddlestun (1976), Schmidt and 
Clark (1980), Huddlestun (1984), Schmidt (1984), Meeder (1987), Huddlestun 
(1988), Scott (1988), Lyons (1991), Missemer (1992), and Scott (1997). 

Pleistocene Stratigraphy 

Three Pleistocene formations in Florida have records of echinoderms, 
including the Anastasia Formation, Satilla Formation and Bermont Formation 
(Figure 2-1). Just as with the older fossiliferous units in the state, stratigraphic 
definitions and boundaries are under debate for the Pleistocene formations. The 
lithology of these units ranges from dominantly quartz sand with limited fossils in 
the Satilla Formation (Huddlestun, 1988), to interbedded quartz sands and 
coquinas in the Anastasia Formation (Scott, 1991), to a shell-rich, unconsolidated 
sandy marl in the Bermont Formation (Dubar, 1962). The thickness of the 


18 


Pleistocene formations ranges from less than one m in outcrop to nearly 38 m in 
the subsurface. Further details regarding stratigraphic nomenclature history, 
lithology, thickness of specific units, and fossil content is present in a variety of 
publications. Selected examples of relevant work (listed chronologically) include: 
Veatch and Stephenson (1911), Sellards (1912), Chamberlin (1917), Cooke 
(1926), Cooke and Mossom (1929), Cooke (1943), Parker and Cooke (1944), 
Cooke (1945), Parker et al. (1955), Dubar (1962), Vokes (1963), Olsson and 
Petit (1964), Olsson (1968), Brooks (1974), Dubar (1974), Blackwelder (1981), 
Kussel and Jones (1986), Huddlestun (1988), Scott (1988), Lyons (1991), Scott 
(1991), and Scott (1997). 


CHAPTER 3 

SYSTEMATIC PALEONTOLOGY OF FLORIDA ECHINODERMS 

Introduction 

The fossil echinoderms of Florida belong to four groups (classes), 
including the echinoids, asteroids, ophiuroids, and crinoids. Whereas fossil 
echinoids are diverse and abundant throughout the Cenozoic of Florida, a limited 
published record or description exists for ail other groups. In fact, until Jones and 
Portell (1989) described the occurrence of Heliaster microbrachius in 
southwestern Florida, most published records of asteroids referred only to the 
presence of unidentified ossicles within several formations. A similar pattern is 
true for the crinoids of Florida, with only one detailed discussion (Oyen, 1995) of 
disarticulated comatulid crinoid ossicles of Himerometra bassleri and a second 
unidentified species. Even less is known about the species and distribution of 
ophiuroids within the state, with the best examples of fossil ophiuroids published 
by Ivany et al. (1990). No compilation of all fossil echinoderms from Florida 
exists, even at class and lower taxonomic levels. This report not only adds 
significant new information to the taxonomic database, but also provides a 
comprehensive, single reference source for biostratigraphers studying such 
fossils in Florida stratigraphic units. 


19 


20 

Though it is not the goal of this dissertation to formally describe all new 
taxa of echinoderms presented herein, it is warranted to provide the taxonomic 
setting for the various groups of fossils included in the discussion. I have chosen 
to leave any formal species descriptions for future work. Herein I provide 
preliminary descriptions of all fossil echinoderms, collected by me or other 
researchers and private collectors, that are new to the biostratigraphic and 
taxonomic realms of the Cenozoic fossil record in Florida. However, I also have 
included formal generic and specific descriptions for all echinoderm taxa that 
have been identified and reviewed in a professional forum. Along with such 
descriptions, I include remarks regarding my interpretation of the taxonomic 
identifications that I have made and the justification for such, as well as how 
these newly reported echinoderms affect the overall biostratigraphic record and 
species diversity estimates for each epoch associated with the fauna. Finally, my 
descriptions of these fossils include terms stating the echinoids are small, 
medium, or large in size. In general, these size terms relate to a test length of 
approximately less than four cm in the small category, less than about eight cm 
in the medium size category, while those echinoid fossils larger than eight cm are 
considered large. 

Each of the four echinoderm class-level sections are presented in a 
phylogenetic context rather than a stratigraphic context, and all formal 
descriptions are limited to genus and species taxonomic levels. However, due to 
the large number of species associated with the Florida echinoids, taxonomic 
information is presented in a phylogenetic context with additional subdivision by 


21 


epoch. Only limited discussion of these taxa is included with regard to diversity 
and taxonomic implications of these results. Detailed examination of diversity 
patterns, changes in the patterns as a result of these new data, biases 
associated with such patterns, and relationship to echinoderm patterns from the 
western Atlantic, Caribbean, and Gulf of Mexico are presented in Chapter 4 of 
this dissertation. 


Class Echinoidea Fossils 

Eocene Echinoids 

Class Echinoidea Leske, 1778 
Order Cidaroida Claus, 1880 
Family Cidaridae Gray, 1825 
Genus Phvllacanthus Brandt, 1835 

Description: Test spherical or low, usually flattened above, sides arched. 
Areoles well separated, central part elevated, carrying prominent, non-crenulate 
primary tubercle. Madreporite conspicuously larger than other genital plates, 
encroaching on small periproct. Scrobicular tubercles conspicuously larger than 
other secondaries, usually with distinct elevation on side toward areole. Pores 
conjugate, but with wall elevated aborally. Primary spines cylindrical, thick, 
robust, with fine granules arranged in regular longitudinal series on shaft; cortex 
thick; primary radial lamellae (as seen in transverse section) arising in fanlike 
clusters from projecting portions of medulla. Secondary spines broad, flat, 
squamiform, closely adpressed. Globiferous pedicellariae without end-tooth. 

Florida species; P. mortoni (Conrad, 1850). 

Comments: The species occurs throughout the Ocala Limestone. 


22 


Phvllacanthus mortoni (Conrad. 1850) 

(Figure 3-1, A-C) 

Material examined: UF 66913 (figured test), UF 3270 (1136 radicles), 

UF 12993 (test with spines), UF 68683 (partial test). 

Description: Test large. Apical system subcircular, larger than 
peristome. Ambulacra nearly uniform in width, ribbonlike; poriferous zones about 
twice as wide as the interporiferous zones; pores round or oval, conjugate; 
zygopores transverse, separated by a ridge; interporiferous zones granular. 
Interambulacra composed of about eight to ten tiers of plates, the median area 
somewhat sunken; plates wider than high; tubercles smooth, perforated; high; 
granules arranged in transverse rows separated by a groove. Peristome 
pentagonal, the angles truncated at the ambulacra. Spines decorated with 
longitudinal rows of spinelets. On some the spinelets are uniform in size; others 
have longer thornlike spinelets at regular intervals. 

Remarks: Few nearly complete fossils of this species have been col- 
lected, but the preservation of test morphology normally remains good although 
fragmentation may have occurred. The species is relatively easy to identify, 
even when radicles are the only skeletal component present at a given locality. 


Order Diadematoida Duncan, 1889 
(Figure 3-1, D-E) 

Florida species: Family, genus, and species undetermined. 

Comments: The specimens were collected from the Upper Ocala 
Limestone in northern Florida and consist of one incomplete test (unavailable for 


23 

description) and several lantern components (figured). These fossils are 
important because they represent both a new stratigraphic record and, with high 
probability, a new taxonomic record for the state. 

Material examined; UF 32929 (figured hemi-pyramids of lantern). 

Order Phymosomatoida Mortensen, 1904 
Family Phymosomatidae Pomel, 1883 
Genus Dixieus (Cooke, 1941) 

Description: Test low-arched above, rounded below. Apical system 
small, pentagonal, equilateral, composed of five genital plates and five ocular 
plates, the anterior and right anterior ocular plates nearly or quite reaching the 
periproct, the other ocular plates widely insert. Periproct central, comparatively 
large. Peristome large, circular, deeply notched, depressed. Ambulacra evenly 
expanded to the ambitus, where they are more than half as wide as the 
interambulacral areas; plates somewhat wider than high, each bearing one 
central, primary, crenulated, imperforate tubercle; poriferous zones above the 
ambitus consisting of two vertical rows of zygopores, about 12 zygopores to each 
compound plate, becoming somewhat disordered near the ambitus, and 
changing below the ambitus to one row of open, connected arcs, one arc of 
about six zygopores to each compound plate, breaking into two diagonal rows of 
three near the peristome. Interambulacral plates about as high as the 
ambulacral but somewhat wider, each bearing one central, imperforate, 
crenulated primary tubercle. Tubercles evenly graded. 

Florida species: D. dixie (Cooke, 1941). 


24 


Comments; The species occurs in the Upper Ocala Limestone in north- 
central Florida. 

Dixieus dixie (Cooke, 1941) 

(Figure 3-1, F-H) 

Material examined: UF 5467 (figured test), UF 66559 (figured test), UF 
5342 (test), UF 5788 (test). 

Description: Test moderately large, circular; upper surface gently 
arched; lower surface evenly rounded, concave near the peristome. Oculogenital 
ring monocyclic; madreporite large, tumid; other genital plates and ocular plates 
with sparsely scattered granules. Ambulacral areas as wide as interambulacral 
areas; pore pairs on upper surface biserial, nearly straight, 10 to 12 pairs on 
each plate, pore pairs of the two series alternating in position; pore pairs on lower 
surface in uniserial arcs, four or five pairs on each plate. Primary tubercles large, 
imperforate, crenulated, two rows on ambulacral areas and two on 
interambulacral areas. Miliary tubercles mammillated, near edges of plates on 
both ambulacral and interambulacral areas. Peristome large, circular; gill slits 
deeper than wide; peristomial edge of ambulacra bilobate. 

Order Oligopygoida Kier, 1967 
Family Oligopygidae Duncan, 1889 
Genus Oliqopyqus de Loriol, 1887 

Description; Slightly concave orally, deep depression around peristome; 
anterior petal usually longest, pores subequal, conjugate; apical system 
subcentral; periproct inframarginal; tubercles imperforate, noncrenulate. 

Florida species: Three species are present, including O. wetherbvi de 
Loriol, 1887, O. haldemani (Conrad, 1850), and O. phelani Kier. 1967. 


25 


Comments: Species of Oliqopyqus are abundant in the Eocene; O. 
phelani . (Lower Ocala Limestone); O. haldemani and O. wetherbyi (Upper Ocala 
Limestone). 

Oliqopyqus phelani Kier, 1 967 
(Figure 3-1, l-J) 

Material examined: UF 18017 (figured test), UF 1645 (test), UF 5854 (28 
tests), UF 12551 (test), UF 38222 (150 tests). 

Description; Test small, elongate; width 83 to 88 percent of length, 
length-width ratio quite constant; in smaller specimens marginal outline oyal, in 
large specimens subpentagonal, pointed anteriorly, blunted posteriorly with 
greatest width anterior; greatest height commonly at apical system, in some 
specimens anterior; height quite yariable, ranging from 42 to 60 percent of 
length; adapical surface slightly conyex, sides smoothly curying, adoral surface 
lacking deep peristomal sulcus, only depressed immediately around peristome 
opening. Apical system central to slightly posterior; monobasal, madreporite 
strongly inflated, seyeral tubercles on madreporite; ocular plates small; four 
genital pores, anterior pair closer together than posterior, pores large in some 
specimens small in others. Petals well deyeloped, open in some specimens, 
straight, slightly closing in others; interporiferous zone widest in petal III where 
almost twice as wide as single poriferous zone; in other petals interporiferous 
zone slightly wider than poriferous zone; petal III longest with four to nine more 
pore pairs in single poriferous zone than petals II or IV, four to seyen more than 
petals V or I; pores strongly conjugate, in sutures between plates. Beyond 
petals, ambulacral plates single pored; at extremity of petal, pores yery 


26 


numerous in many included ambulacral plates and demiplates; at ambitus pores 
most crowded in double series in each half ambulacrum with continuous column 
of demiplates separating primary plates from adradial suture; a few included 
plates inserted between primaries and demiplates; included and demiplates near 
adradial border, plates thin not extending through test; nearing peristome primary 
plates extend to adradial suture, no included plates, one demiplate for each 
primary; buccal pores difficult to see. Two columns in interambulacral areas 
except at peristome, where column terminating in single plate. Peristome slightly 
anterior, central, or slightly posterior, opening slightly wider than high, curved 
anteriorly, slightly pointed posteriorly; not deep in sulcus, test only depressed in 
area immediately around opening. Periproct small, slightly wider than high; 
located between 54 and 73 percent of the distance from center of peristome to 
posterior margin. Test covered with small, irregularly arranged tubercles; 
scrobicules deep with vertical side; boss large, two-thirds diameter of scrobicule, 
extending upward as high as surrounding surface of test; crenulated; mamelon 
small, extending in height above surface of test, perforated; crenulations and 
mamelon present only in well preserved specimens; small secondary tubercles 
scattered over area between tubercles. 

Oliqopyqus haldemani (Conrad. 1850) 

(Figure 3-1, K-L) 

Material examined: UF 47257 (figured test), UF 38585 (83 tests), UF 
46715 (35 tests), UF 47956 (13 tests). 


27 

Description; Horizontal outline broadly ovate, nearly circular when 
juvenile; upper surface nearly flat in front, slightly turned down behind and slop- 
ing toward the peristome; margin broadly rounded. Apical system central, tumid, 
monobasal, with four genital pores. Petals short, extending little more than 
halfway to the margin, the anterior the longest, expanding distally and open at 
the tips; poriferous zones narrower than the interporiferous; pores circular, 
conjugate; plates narrow. Extrapetaliferous parts of ambulacra expanding to the 
margin; plates wider. Interambulacra thickened around the peristome internally. 
Peristome central, oval, deeply sunken in a transversely elongated pit whose 
anterior side stands nearly vertical and whose posterior side slopes less steeply 
and extends about halfway to the margin. Auricles erect; prongs erect, separate, 
far apart. Jaws present. Periproct small, circular, submarginal. Tubercles small, 
sunken. 

Oliqopyqus wetherbyi de Loriol, 1887 
(Figure 3-1, M-N) 

Material examined; UF 17756 (figured test), UF 17770 (test), UF 40052 
(18 tests), UF 40607 (21 tests), UF 47955 (three tests). 

Description; Horizontal outline broadly ovate, nearly circular when 
young; upper surface moderately tumid; lower surface nearly flat in front, slightly 
turned down behind and sloping toward the peristome; margin broadly rounded. 
Apical system central, tumid, monobasal, with four genital pores. Petals short, 
extending little more than halfway to the margin, the anterior the longest, 
expaning distally and open at the tips; poriferous zones narrower than the 
interoriferous; pores circular, conjugate; plates narrow. Extrapetaliferous parts of 


28 


ambulacra expanding to the margin; plates wider. Interambulacra thickened 
around the peristome internally. Peristome central, oval, deeply sunken in a 
transversely elongate pit whose anterior side stands nearly vertical and whose 
posterior side slopes less steeply and extends about halfway to the margin. 
Auricles erect; prongs erect, separate, far apart. Jaws present. Periproct small, 
circular, submarginal. Tubercles small, sunken. 

Family uncertain 

Genus Amblypygus L. Agassiz, 1840 

Description: Circular to ovate, low-arched to high subconical, flattened 
orally, margin tumid; ambulacra petaloid, pores conjugate, outer pore elongate, 
pores small adorally; apical system apparently tetrabasal; peristome sunken, 
subrounded to oblique; periproct large, pyriform, inframarginal; tubercles 
perforate, crenulate; no evidence of girdle or lantern in adult. 

Florida species: A. americanus Michelin, 1858. 

Comments: The species is present in the Upper Ocala Limestone 

Amblypygus americanus Michelin, 1858 
(Figure 3-2, A-B) 

Material examined: UF 67090 (figured test), UF 648 (test), UF 5254 
(test), UF 22133 (2 tests). 

Description: Test large; horizontal outline circular; upper surface more or 
less inflated; lower surface gently rounded, concave around the peristome and 
less so around the periproct; margin broadly rounded. Apical system central. 


29 


with four genital pores; a large central madreporite fills in the spaces beside the 
much smaller genital plates; ocular plates small. Ambulacra narrow, continuous, 
expanded on upper side into petals with round inner pores and elongated outer 
pores; petals extending to margin, inner pores forming a straight line, outer sides 
of petals slightly arched. Peristome subtriangular, oblique, wider than long, 
without floscelle. Periproct on the lower surface, large longer than wide, nearer 
the peristome than the margin. Tubercles small, sunken; intermediate spaces 
and madreporite granulated. 

Order Clypeasteroida A. Agassiz, 1872 
Family Fibulariidae Gray, 1855 
Genus Fibularia Lamarck, 1816 

Description; Test ovate, inflated; periproct close to peristome; 
hydropores in groove; no internal supports; five large periproctal plates; buccal 
membrane naked; no calcareous disc in tube feet. 

Florida species: F. vauqhani (Twitchell, 1915). 

Comments; The species is present in the Lower Ocala Limestone, and 
also represents a new stratigraphic record from the Upper Ocala Limestone. 

Fibularia vauqhani (Twitchell. 1915) 

(Figure 3-3, A-B) 

Material examined: UF 38211 (figured test), UF 18014 (five tests), UF 
27521 (500 tests), UF 38208 (100 tests). 

Description; Test very small, thick walled, elongate egg shaped in 
general form, elongate subelliptical to elongate subovate in marginal outline. 


30 


somewhat pointed anteriorly, about twice as long as broad. Aboral surface is 
high, with height equal to width, and flattened longitudinally; adoral surface also 
flattened somewhat along the longitudinal median area and slightly concave 
around the peristome. Ambulacral petals rather well defined, relatively short; 
petals wide open at ends; poriferous zones diverge in almost straight lines to the 
ends and consist of small round pores in pairs not distinctly conjugated. 
Peristome is relatively large, central, slightly depressed below the surface. 
Periproct is very small, about one-third the diameter of the peristome and very 
close to the peristome. Apical system anteriorly eccentric, with four medium- 
sized genital pores. 


Family Neolaganidae Durham, 1954 
Genus Neolaqanum Durham, 1954 

Description: Small to medium-sized; petals nearly closed, length 0.7 of 
radius; plates within petals in dyads and triads; four genital pores; hydropores in 
branching groove; periproct oral, about 0.25 distance from margin; four or five 
ambulacral and three or four interambulacral coronal plates per column on oral 
surface. 

Florida species; Two total, including N. dalli (Twitchell, 1915) and J)J. 
durhami Cooke, 1959. 

Comments: Both taxa are present in only one formation each. N. dalli is 
diagnostic of the Avon Park Formation while N. durhami is present in the Upper 


Ocala Limestone. 


Neolaqanum dalli (Twitchell, 1915) 

(Figure 3-3, C-D) 

Material examined: UF 104422 (figured test), UF 18385 (81 tests), UF 


31 


28184 (23 tests). 

Description: Test small, subpentagonal in marginal outline. Entire form 
greatly depressed, subdiscoidal, the upper surface nearly parallel with the lower. 
Test margin notably thickened, slightly less so at the middle of the posterior end. 
Lower surface flat or nearly so. Apex subcentral or very slightly eccentric 
anteriorly, though it is but slightly higher than the margin of the test. There is no 
distinct concave ring on the upper surface, but the poriferous zones of the 
ambulacral petals are slightly depressed below the general surface. Ambulacral 
petals subovate in outline, broad, extending nearly two-thirds the way to the 
margin, rounded, blunt, and closed at the ends. Poriferous zones very wide, as 
wide as the interporiferous areas, slightly depressed below the general surface; 
inner row of pores round, outer row slit-like, pairs of pores conjugated by very 
narrow grooves. Interporiferous areas narrow, standing in relief by reason of the 
depression of the poriferous zones. The whole surface of the test, including the 
interporiferous areas, covered with small tubercles, set in deep scrobicules, 
which are larger on the adoral surface. Apical system subcentral or slightly 
eccentric anteriorly, coincident with the apex. Four large genital pores, of which 
the anterior pair are nearer together than the posterior pair. Madreporite tumid. 
Peristome central or subcentral in position. Periproct small, circular, about 
midway between the margin and the peristome. 


Neolaqanum durhami Cooke, 1959 
(Figure 3-3, E-F) 


32 


Material examined; UF 13026 (figured test), UF 13041 (test), UF 30599 
(75 tests), UF 38581 (100 tests). 

Description: Fiorizontal outline broadly oval to subdecagonal; upper side 
slightly tumid at the apex, depressed sub-marginally; margin usually thick, 
rounded: lower side flat. Apical system nearly central; four genital pores rather 
far apart; hydropores in crooked grooves. Petals long, extending nearly three- 
fourths the radius, equal, spatulate; poriferous zones nearly as wide as the 
interporiferous zones, nearly closed distally, open at the apex; inner pores 
circular or oval, outer pores elongated; plates compound. Peristome small 
central, pentagonal; teeth usually preserved; five pairs of large buccal pores. 
Ambulacral grooves straight, extending nearly halfway to the margin. Periproct 
small, circular, about three-fourths the radius from the peristome. Tubercles 
small, sunken, numerous. Internal concentric passageways near the margin. 

Genus Durhamella Kier, 1968 

Description; Test small to medium in size, low, with flat adoral surface. 
Plates of adapical surface may or may not be tumid, with sutures depressed. 

The apical system has five genital pores, and the pores occur within or without 
the fused genital plates. The hydropore opens in one or two slits. The petals are 
wide near the apical system and extend approximately one-half the distance from 
the apical system to the margin. The pores are conjugate, but the outer pore is 
not in a slit, only elongated transversely. Pseudocompound plates are present in 


33 


the petals with approximately six to eight in each petal. The accessory pores 
occur along the transverse sutures of the ambulacral plates adapically, but 
adorally throughout the basicoronal and first coronal ambulacral plates. The 
interambulacra terminate at the apical system, with a single quadrangular plate. 
Adorally, the basicoronal plates have a circular to subpentagonal outline, with a 
single plate in each interambulacrum, double plates in each ambulacrum. The 
first coronal interambulacral plates are high, extending beyond the first coronal 
ambulacral plates. The periproct is inframarginal, and the interior supports are 

concentric. No food grooves are present. 

Florida species; Two species are present in Florida, including D. 
floridana (Twitchell, 1915) and D. ocalana (Cooke, 1942). 

Comments: Both species are present in the Upper Ocala Limestone. 

Durhamella floridana (Twitchell, 1915) 

(Figure 3-3, G-H) 

Material examined: UF 3356 (figured test), UF 3358 (test), UF 48135 
(two tests). 

Description: Small test, almost regularly ova! in marginal outline. Entire 
form is greatly depressed; subdiscoidal; upper surface almost parallel with the 
lower; the apical region slightly tumid, the tumidity involving the larger part of the 
petals; the region around the ends of the petals concave; the margin notably 
thickened, slightly more so anteriorly than posteriorly. Lower surface flat, or 
nearly so. Apex slightly eccentric anteriorly, at the summit of the central tumid 
area, which rises but very slightly above the height of the margin. Ambulacral 


34 


petals subelliptical, somewhat pointed and closed at the ends; very short, 
extending only about halfway to the margin, subequal in length. Poriferous 
zones very narrow, much narrower than the interporiferous areas, the proximal 
ends poorly defined, inner row of pores round, outer row slit-like, pairs of pores 
conjugated. Entire surface of test covered with small tubercles set in deep 
scrobicules, which are somewhat larger on the lower surface. Apical system 
slightly eccentric anteriorly, coincident with the apex. Four large genital pores, 
the anterior pair being set closer together than the posterior pair. Poriferous 
zones do not come together at their proximal ends, and the perforations of the 
radial plates not visible. Peristome small, slightly eccentric anteriorly, 
subpentagonal. Ambulacral grooves inconspicuous. Periproct small, about half 
the diameter of the peristome, circular, located about one-third the way from the 
margin to the peristome. 

Durhamella ocalana (Cooke. 1942) 

(Figure 3-3, l-J) 

Material examined: UF 3341 (figured test), UF 38600 (100 tests), UF 
65706 (14 tests). 

Description: Outline subpentagonal to oval; margin thick; submargin 
depressed; petaloid region tumid; lower surface flat. Individual plates in the area 
between the thick rim and the petals tumid. Apical system slightly eccentric 
anteriorly; five genital pores. Petals short, extending about halfway to the 
margin; nearly closed; poriferous zones much narrower than interporiferous 
zones, widely separated at the proximal ends. Peristome very small, pentagonal, 


35 


central. Periproct about one-quarter way from the margin to the peristome. 
Tubercles fairly large, of uniform size; more widely scattered on the lower than on 
the upper surface. 


Genus Weisbordella Durham, 1954 

Description: Like Neolaqanum but with larger periproct and without 
groove for hydropores; oral surface slightly concave. 

Florida species; Two species are present in Florida, including 
Weisbordella cubae (Weisbord, 1 934) and W. iohnsoni (Twitchell, 1915). 

Comments. Both species are present in the Upper Ocala Limestone. 

Weisbordella cubae (Weisbord. 1934) 

(Figure 3-3, K-L) 

Material examined: UF 5846 (figured test), UF 5839 (test), UF 41280 
(162 tests), UF 47164 (17 tests), UF 64432 (3 tests). 

Description: Test very small, thin, subovate, longer than broad, slightly 
thicker at the anterior margin than at the posterior. Dorsal face somewhat 
conical, low, rising somewhat above the level of the margins, with a subdued 
ridge in the posterior interradium extending from near the disc to the margin. 
Peristome subcentral, perhaps a bit eccentric anteriorly. Periproct flush, 
subpentagonal, a short distance from the posterior margin, in the antero-posterior 
plane of symmetry. Apical disc slightly eccentric anteriorly. Ambulacral petals 
subequal, petaloid, broad at the disc, pointed and closed distally, not quite 
reaching the margin. Interporiferous zone moderately convex, about twice as 


36 


wide as the poriferous zone which is virtually flush, and arrayed with oblique, 
conjugate pore pairs, the pores of a pair not distinct but seemingly subequal. 

The test is ornamented with rather small, lightly scrobiculate tubercles, less 
numerous and fainter dorsally than ventrally. At the apex are three or four 
tubercles somewhat larger than average, though impressed about the same. In 
addition to these are four tubercles larger than any of the others and more 
prominently scrobiculate, two of which are in interambulacrum 2, and one each in 
interambulacra 3 and 4, situated near the distal ends of the petals. 

Weisbordella iohnsoni (Twitchell, 1915) 

(Figure 3-3, M-N) 

Material examined; UF 47957 (figured test), UF 5807 (test), UF 12985 
(test), UF 37553 (test). 

Description: Test moderate in size and almost regularly oval in marginal 
outline. Upper surface moderately elevated centrally; height about one-third of 
the width; the tumid area extending to the ends of the petals; the submarginal 
area about equal in thickness to the margin, which is slightly undulating, very 
thick, high and rounded, thicker and higher than in related forms, slightly thinner 
at the middle of the posterior end than elsewhere. Lower surface decidedly 
concave; the concavity reaching nearly to the margin and near the peristome 
being about equal to one-half the height of the test. Apex subcentral. Posterior 
petals lanceolate; the anterior three subelliptical, all of them pointed and closed 
at the ends, extending two-thirds or more of the way to the margin; anterior pair 
slightly shorter than the rest. Poriferous zones very narrow, much narrower than 


37 


the slightly tumid interporiferous areas, sometimes irregular, inner ends poorly 
defined, inner row of pores round, outer row slit-like, pairs of pores conjugated. 
Whole surface of test covered with rather conspicuous small tubercles which are 
larger on lower surface. Apical system subcentral, with four large genital pores. 
Peristome small, subpentagonal, subcentral; ambulacral grooves poorly defined. 
Periproct small, subcircular to subpentagonal, about one-third the way from the 
margin to the peristome. 


Genus Wvthella Durham, 1954 

Description: Similar to Cubanaster but larger, margin thinner, petals 
raised and interambulacral areas widened midway on oral surface; also similar to 
Neorumphia but interambulacra much narrower at ambitus. 

Florida species: W. eldridqei (Twitchell, 1915). 

Comments: The species is present in the Upper Ocala Limestone. 

Wvthella eldridqei (Twitchell. 1915) 

(Figure 3-3, 0-P) 

Material examined: UF 5803 (figured test), UF 39541 (46 tests), UF 
46933 (25 tests), UF 48492 (11 tests). 

Description: Test large, subpentagonal to subdecagonal in marginal 
outline, longitudinally elongate, truncated at the anterior and posterior ends, more 
or less undulating along the sides. Whole form greatly depressed, margin thin 
but thicker than slightly concave submarginal area, petaloidal region tumid. Apex 
and apical system subcentral. Four large genital pores are present, with the 


38 


anterior pair set closer together than the posterior pair. Whole test is closely set 
with very small tubercles, among which are scattered at irregular distances some 
larger ones in deep scrobicules. Lower surface flat. Ambulacral petals long, 
elongate elliptical, extending two-thirds the way to the margin, pointed and closed 
at the ends; pairs of pores conjugated by very narrow more or less wavy 
grooves. Ambulacral areas very wide at margin, narrowing rapidly to ends of 
petals. Peristome moderate in size, subcentral, subpentagonal to subelliptical, 
transversely elongate. Ambulacral grooves apparently simple and straight, each 
groove having a fine line on both sides which rapidly diverge from the main 
groove. Periproct relatively large, suboval, longitudinally elongate, one-fourth the 
way from the margin to the peristome. 

Family Protoscutellidae Durham, 1955 
Genus Protoscutella Stefanini. 1924 

Description: Cooke describes the test as low, ambitus thin, usually with 
posterior periproctal notch; petals equal, length about half of radius; periproct 
submarginal, between third and fourth coronal plates; food grooves simple, 
nonbranched; posterior interambulacrum discontinuous; six or seven ambulacral 
and three to five interambulacral coronal plates on oral surface. 

Florida species; P. pentaqonium Cooke, 1942. 

Comments; This species is problematic with respect to biostratigraphy. 
Cooke (1942) reported this echinoid from a water (?) well drilled in Washington 
County, Florida, and stated the age of the rock as middle Eocene in age, possibly 
representing the Lisbon Formation. I have not been able to locate the specimen 


39 

he referred to in his paper, and therefore I can only consider the formation in 
which the fossil is found to be uncertain. 

Protoscutella pentaqonium Cooke, 1942 
Material examined; No UF specimens were available for examination. 
Description: Test subpentagonal, tumid medially, margin thin, slightly 
bent down; covered with small tubercles. Apical system central; five genital 
pores. Petals equal in length, lanceolate, open at the ends, extending halfway to 
the margin; interporiferous zones about equal in width to poriferous zones; 
anterior petal swollen at the apical end, somewhat wider than the others. Lower 
surface flat except for the turning down of the margin; ambulacral furrows 
straight, extending almost to the margin. Peristome small, central, round. 
Periproct round, one-fourth the way from the margin to the peristome; connected 
with the margin by a shallow furrow. 

Genus Periarchus Conrad, 1866 

Description: Test raised apically, ambitus thin; petals open, slender, 
length slightly over half of radius, anterior longest; periproct oral, nearly half 
distance from peristome, between first pair coronal plates; food grooves bifurcate 
about midway on oral surface; all interambulacra continuous; usually seven 
ambulacral and four or five interambulacral coronal plates on oral surface. 

Florida species; P. lyelli floridanus Fischer. 1951. 

Comments: This species is present in the Lower Ocala Limestone. 


40 

Periarchus lyelli floridanus Fischer. 1951 
(Figure 3-4, A-B) 

Material examined; UF 17913 (figured test mold), UF 12795 (figured test 
mold), UF 1187 (test), UF 12754 (test mold), UF 45565 (two test fragments). 

Description; Test large, flat, subcircular, with somewhat wavy or poly- 
gonal margin. Petaloid area slightly tumid, surrounded by a terrace from which a 
gently inclined bevel leads to the sharp margin. Apical system central or nearly 
so, apex at centrally located madreporite or at base of anterior petal. Five genital 
pores. Anterior petal longest, posterior petals intermediate, antero-lateral petals 
shortest. Longest petals reach nearly halfway to margin. Petals are broadly 
lanceolate, reaching maximum diameter near the midpoint. Their outer margins 
are more or less regular convex arcs except distally where in contact with the first 
pair of non-petaloid ambulacral plates; here petals are abruptly terminated by 
concave margins. Interporiferous zones are broadly lanceolate at proximal end, 
widen gently to one-half to three-quarters of their length, then taper gradually to 
the narrow, open, distal end. Poriferous zones begin narrow, widen for half their 
length, then maintain their width until near the end, where they suddenly taper to 
produce the concave margins. Inner pores are slightly elliptical; outer ones form 
long, narrow slots. Oral surface flat. Peristome small; position subcentral, 
variable. Periproct very small, nearer to peristome than to margin. Five actinal 
grooves bifurcate at angle of 30° to 35°, about halfway to margin. At margin 
interambulacral areas are slightly larger than the ambulacral areas. The tumid 
central region houses a large Aristotle’s lantern; remainder of test shows a 


41 

complex internal septation. Each of the peripheral plates carries a system of 
septa. Inner plates carry a few low septa or ridges radiating out from peristome. 

Genus Mortonella Pomel, 1883 

Description: Like Periarchus but test thick, margin rounded, petals 
broader, and periproct midway on oral surface. 

Florida species: Two taxa are present, including M. quinguefaria (Say, 
1825) and M. quinguefaria kewi Cooke, 1942). 

Comments: These echinoids were reported from the Crystal River 
Formation (Upper Ocala Limestone), although the formation they were collected 
from is still questionable. Cooke (1959, p. 44) stated he did not believe this 
genus is valid and, at best, should be a subgenus of Periarchus . 

Mortonella quinguefaria (Say. 1825) 

(Figure 3-5, A-B) 

Material examined: UF 2202a & b (two tests figured), UF 2203 (12 
tests), UF 2204 (7 tests). 

Description: Horizontal outline circular; upper surface slightly tumid 
centrally, submargin flat; margin generally rather thick or beveled, oral side flat. 
Apical system central, having a large central madreporite and five genital pores. 
Petals wide, nearly equal in length, extending three-quarters of distance to 
margin, outer edges convex; poriferous zones wider than the interporiferous 
zones, closed at the apical ends, open distally; inner pores circular, in nearly 
straight lines; outer pores elongated; pores conjugate. Peristome small, circular. 


42 


central; food grooves extending halfway to margin, where the grooves bifurcate; 
each branch of the grooves has an outward branching extending nearly at right 
angles to it before curving to the margin; grooves punctate. Periproct smaller 
than the peristome, circular, nearly midway between the peristome and margin. 
Sunken tubercles cover entire surface, including the apical system. 

Mortonella quinquefaria kewi (Cooke, 1 942) 

(Figure 3-5, C-D) 

Material examined: UF 5275 (figured test). 

Description: Test circular, margin weakly fluted, thin, covered with small 
tubercles. Apical region tumid; five genital pores at the same distance from the 
center as the apical ends of the petals. Petals nearly closed at the apical ends; 
wider open at the outer ends; interporiferous zones somewhat narrower than 
poriferous zones, nearly straight, widening slightly distally; inner pores round, 
connected by grooves with the slot-shaped outer pores; petals extending three- 
fifths the way to the margin. Petals encircled by a tumid ring, which gives the 
submarginal area a beveled appearance. Base flat; ambulacral furrows 
branching about halfway to the margin; buccal tubes extending to the bifurcation. 
Peristome nearly circular, sunken, about twice as large as the periproct. 

Periproct small; midway between the peristome and the margin. 


43 


Order Cassiduloida Claus, 1880 
Family Echinolampadidae Gray, 1851 
Genus Echinolampas Gray, 1825 

Description; Medium-sized to large, usually inflated; apical system 
monobasal; poriferous zones usually unequal, wide interporiferous zones. 

Florida species: E. tanvpetalis Harper and Shaak, 1974. 

Comments: This species is present in the Upper Ocala Limestone. 

Echinolampas tanvpetalis Harper and Shaak, 1974 
(Figure 3-5, E-F) 

Material examined: UF 5385 (figured test), UF 2283 (test), UF 3378 
(test), UF 3900 (test). 

Description; Test large, ovate to roundly subpentagonal; greatest height 
near center; anterior margin broadly rounded; posterior margin broadly rounded 
adapically, somewhat sloping adorally; adoral surface depressed around 
peristome. Apical system anterior of center, and approximately 44 percent of test 
length from anterior margin; monobasal, with four genital pores. Petals very 
long, extending almost to margin, broad, with interporiferous zones greater than 
four times width of poriferous zones; pores conjugate, nearly perpendicular to 
petal axis, outer pores transversely elongated, inner pores round; poriferous 
zones of unequal length, more pore pairs in right zone of petal III, posterior zones 
of petals II and IV, and outer zones of petals I and V; petals I, III, and V straight, 
petals II and IV flexed slightly toward anterior; petals with no tendency to close 
distally. Periproct inframarginal, transverse, large roundly triangular. Peristome 
anterior of center, distance from anterior margin approximately 42 percent of test 


44 


length; transverse, large, roundly triangular to subpentagonal. Bourrelets poorly 
developed; phyllodes well developed, narrowing toward peristome, single-pored 
with many pores arranged in two series in each half ambulacrum: in ambulacrum 
III, two series in each half, with eight pores in outer series, three to four in inner 
series; in others, eight pores in outer series, four to six in inner series; buccal 
pores present. Tubercles small, somewhat depressed, more numerous and 
smaller on adoral surface. 

Family Cassidulidae L. Agassiz and Desor, 1847 
Genus Rhvncholampas A. Agassiz, 1869 

Description: Small to large, oral surface flat; apical system monobasal or 
tetrabasal; periproct supramarginal to marginal, longitudinal or transverse; 
peristome transverse; petals broad, usually equal, commonly inconspicuous, 
ambulacral plates double-pored in pre-Senonian species; tubercles much larger 
adorally, naked zone in interambulacrum 5 adorally. 

Florida species: A total of six taxa (two of which being subspecies) are 
present in Florida, including R. con rad i (Conrad, 1850), R. conradi Ivelli (Conrad, 
1850), R. ericsoni (Fischer, 1951), R. qeorqiensis (Twitchell, 1915), R. 
qeorqiensis qlobosus (Fischer, 1951), and R. troianus (Cooke, 1942). 

Comments: The taxa in this genus are present throughout the Ocala 
Limestone, with R. conradi , R. conradi Ivelli. and R. troianus preserved in the 
Upper Ocala Limestone and R. ericsoni . R. qeorqiensis . and R. qeorqiensis 
qlobosus part of the Lower Ocala Limestone. 


Rhvncholampas conradi (Conrad, 1850) 

(Figure 3-6, A-C) 

Material examined: UF 39452 (figured test), UF 39453 (13 tests), UF 


45 


64904 (9 tests), UF 65809 (16 tests). 

Description; Florizontal outline oval, usually somewhat compressed 
posterolaterally; upper surface evenly tumid, rostrate above the periproct; lower 
surface usually slightly concave around the peristome; margin broadly rounded. 
Apical system anteriorly eccentric, monobasal, with four genital pores. Petals 
long and rather straight, open distally; inner poriferous zones of paired petals 
longer than the outer; inner pores circular, outer pores oval. Peristome large, 
pentagonal, wider than long, nearer the center than the apical system; phyllodes 
nearly as wide as long; bourrelets elongated, swollen, granulated. Periproct 
terminal, at the top of a narrow vertical truncation, wider than high. Tubercles 
depressed, small on upper surface, larger on lower; covering entire test except a 
narrow pitted band behind the peristome. 

Rhvncholampas conradi lyelli (Conrad, 1 850) 

(Figure 3-6, D-F) 

Material examined; UF 3343 (figured test), UF 3350 (2 tests), UF 47100 
(2 tests). UF 48502 (2 tests). 

Description: This variety is more nearly oval than typical R. conradi , 
lacking the posterolateral constriction that gives R. conradi a pointed 
appearance. It is proportionately somewhat longer and narrower than R. conradi 


carolinensis. 


46 

Rhyncholampas ericsoni (Fischer. 1951) 

(Figured 3-6, G-l) 

Material examined; UF 37661 (figured test), UF 38223 (7 tests), UF 
41270 (8 tests), UF 46635 (test). 

Description: Test outline subpentagonal-subhexagonal, spatulate, wider 
behind than in front. Aboral surface a rather high, inflated cone, uniformly 
covered with small scrobicules. Apex at or just forward of the apical system, 
which is decidedly eccentric toward the anterior. Behind the apex a gentle 
rostrum leads to the upper margin of the periproct. Below the periproct a broad, 
shallow sulcus extends to the margin. Along the sides of the test the inflated 
aboral surface overhangs the sharp margins. Oral surface comparatively flat, 
most prominent at sides, slightly concave around peristome, rises markedly 
toward posterior margin. Three specimens show a slight anterior sulcus. 
Peristome directly below apical system, slightly wider than long, surrounded by a 
large floscelle with prominent bourrelets. Oral surface covered with large, deep 
scrobicules except on anterior and posterior median bands, which are finely 
granulate. Posterior band inflated. Periproct at about half the total height, 
transversely elongate, upper margin gently arched, lower margin more or less 
eccentrically angular. Apical system with four genital pores and central 
madreporite. Petals lanceolate, open distally, nearly equal in length. 
Interporiferous zones in mid-portion two to three times as wide as poriferous 
zones. On posterior petals the anterior pore rows are longer than the posterior 
rows. Pores conjugate; inner ones almost circular, outer ones tapering inward. 


47 

Ridges between pore pairs carry one to three scrobiculate tubercles. The 
number of pores in a row varies from around 40 to 46. 

Rhyncholampas qeorqiensis (Twitchell. 1915) 

(Figure 3-6, J-K) 

Material examined: UF 22535 (figured test), UF 12744 (3 tests). 

Description: Test broadly oval in marginal outline, more obtusely 
rounded posteriorly than anteriorly, and obliquely truncated at the posterior end. 
Upper surface is regularly convex, moderately elevated, in the form of a low, 
rounded ridge above the periproct, sides and anterior end rounded and inflated; 
adoral surface flat, curving upward slightly posteriorly to meet the oblique 
posterior truncation in an angular margin, the angle formed being about 75°. 

Apex is subcentral. Ambulacral areas are narrow, dorsal portions petaloid, the 
petals narrow, partly open at the ends, the posterior petals longer than the 
others, which are nearly equal in length. Poriferous zones are narrow; the inner 
zone of each of the posterior pair of petals shorter than the outer zone; outer row 
of pores slit-like, inner row round; pairs of pores conjugate. Aboral surface of the 
test is closely set with numerous small tubercles which increase in size on the 
adoral surface except along a rather wide median band which is free from 
tubercles and dotted with numerous small pits. Tubercles are set in scrobicules 
that are shallow and irregularly shaped on the aboral surface; but become larger, 
deeper and more regular in form on the adoral surface. Apical system is 
eccentric anteriorly, with four genital pores, of which the anterior pair are nearer 
together than the posterior, and five small perforated radial plates. Peristome is 


48 


eccentric anteriorly, immediately beneath the apical system, pentagonal, 
transversely elongate, with a floscelle of which the phyllodes are rather well 
defined and the bourrelets are large and prominent. Periproct is relatively small, 
about 3 or 4 mm in length, subelliptical to subrhomboidal, transverse; and located 
relatively high up on the posterior surface, at the top of the rather high posterior 
truncation, beneath a rounded, transverse, somewhat protruding expansion of 
the test. 


Rhyncholampas qeorqiensis qlobosus (Fischer. 1951) 

(Figure 3-6, L-0) 

Material examined; UF 32945 (figured test), UF 36435 (figured test), UF 
3342 (2 tests), UF 28182 (5 tests), UF 39094 (test). 

Description: Test medium to large for genus, highly inflated, outline 
nearly circular, profile four-fifths to nearly as high as long. Oral surface 
moderately convex, joining sides at an ill-defined, rounded edge of spatulate 
outline: widest in posterior third. Peristome and apex anteriorly eccentric. 
Peristome pentagonal, transversely elongate, with narrow, beaded bourrelets. 
Periproct at half height, its arched upper margin projecting, lower margin forming 
a broad “V” shape. Test gently rostrate above periproct; a flat band with faint 
median ridge leads from periproct to margin. Sides of test faintly divided into 
similar vertical facets. The sides of the test overhang the oral wall on all sides, 
but much more so in front than in the rear, to produce a forward-leaning appear- 
ance. Top and sides of test covered with small scrobicules. On the margins of 
the oral side these grade into large scrobicules which cover the latter except on 


49 


the anterior and posterior longitudinal median bands, which are finely beaded. 
Apical system with four large genital and five small ocular pores and central 
madreporite. Petals slender, lanceolate, wide open at the ends. On each antero- 
lateral petal the posterior row of pores is longer than the anterior row, whereas 
on each of the posterior petals the anterior row of pores is longer than the 
posterior row. Inner pores round to slightly elliptical; outer pores slightly ovoid. 

Rhvncholampas troianus (Cooke. 1942) 

(Figure 3-6, P-Q) 

Material examined: UP 3747 (figured test), UP 41273 (test), UP 45915 
(test), UP 48497 (12 tests). 

Description: Outline subquadrate, wider posterior than at anterior. 

Upper surface moderately inflated except behind the periproct, where there is a 
broad, shallow sulcus; rostrate above the periproct. Lower surface flat. Margin 
acute. Apical system slightly eccentric anteriorly; four genital pores, five ocular 
pores, madreporite central. Petals lanceolate, of nearly equal length, extending 
somewhat more than halfway to the margin, open at the distal ends; pores round 
or oval; interporiferous zones wider than poriferous zones; outer poriferous zones 
of posterior paired petals longer than the inner. Peristome farther forward than 
the apical system, pentagonal, slightly wider than long. Oral lobes swollen. 
Phyllodes about as long as the diameter of the peristomial opening. Periproct 
supramarginal, transversely elliptical, flush, about one-third the way from the 
margin to the apex. Upper surface finely granulated between small tubercles; 
tubercles on lower surface much larger than on upper, deeply sunken in large 


50 


scrobiculae except near the margin, where they are much smaller. Longitudinal 
median band on base moderately wide and deeply pitted. 

Family Pliolampadidae Kier, 1962 
Genus Eurhodia Haime, 1853 

Description: Medium to large, elongate, low to moderately inflated; 
adorally flattened, apical system monobasal; petals equal, broad, closing distally, 
ambulacral plates beyond petals single pored; periproct supramarginal, 
transverse or longitudinal; peristome higher than wide; bourrelets strongly 
developed; phyllodes broad, single pored, with two series of pores in each half- 
ambulacrum; buccal pores present; tubercles perforate, considerably larger 
adorally than adapically, except for naked and often pitted adoral 
interambulacrum 5. 

Florida species: E. patelliformis (Bouve, 1851). 

Comments: The species is present in the Upper Ocala Limestone. 

Eurhodia patelliformis (Bouve, 1851) 

(Figure 3-7, A-B) 

Material examined: UF 4932 (figured test), UF 3321 (3 tests), UF 41265 
(test), UF 47103 (2 tests), UF 47959 (test). 

Description: Oblong-ovate, and rather pointed posteriorly; horizontal 
outline semicircular in front, narrowly truncated behind. Superior face concavo- 
convex, and forming with the inferior an acute margin. Apical system slightly 
anterior, directly above the periproct, monobasal, four genital pores. Ambulacral 


51 


areas narrow; petals lanceolate, closed at apex, open distally, the anterior pair 
somewhat shorter than the others; poriferous zones of nearly equal length, 
somewhat narrower than the interporiferous; inner pores circular, outer pores 
oval or elongate. Peristome pentagonal, slightly longer than wide; bourrelets 
strong, bulbous. Periproct supramarginal, nearly circular, opening into a shallow 
sulcus, which indents the margin and continues forward into the periproct, 
forming an internal shelf or tube. Upper surface and margin covered with small 
sunken tubercles; tubercles on lower surface much larger, sunken in deep pits; 
posterior interambulacrum on lower surface without tubercles but deeply pitted; 
similar pits surround the anterior phyllode. Test width-length ratio approximately 
0.7 and height-length ratio approximately 0.45. 

Order Spatangoida Claus, 1876 
Family Hemiasteridae Clark, 1917 
Genus Ditremaster Munier-Chalmas. 1885 

Description; Subglobular, with faint frontal sinus; two gonopores; paired 
ambulacra petaloid, posterior pair very short, about 0.3 length of anterior ones. 

Florida species; D. becked (Cooke, 1942). 

Comments; The species is present in the Upper Ocala Limestone. 

Ditremaster becked (Cooke, 1942) 

(Figure 3-7, C-D) 

Material examined; UF 47040 (figured test), UF 4922 (2 tests), UF 5819 


(test), UF 41274 (test). 


52 


Description; Test tumid above and below, margin rounded, highest 
behind the center, sloping steeply forward, vertically truncated behind. Apical 
system behind the center, with two posterior genital pores. Anterior ambulacral 
area deeply depressed halfway to the margin, then becoming shallower, but 
depression still perceptible at the peristome. Petals deeply depressed; anterior 
pair long, with nearly straight margins, slightly curved outwards at the tip; 
posterior pair depressed, much shorter; interporiferous zones about as wide as 
poriferous zones; pores elongate oval, conjugate. Peristome fairly large, curved, 
with a strongly rostrate posterior lip; near the anterior end. Periproct oval, higher 
than wide, high up on the posterior end. Peripetalous fasciole deeply indented 
between the lateral petals. 

Family Schizasteridae Lambert, 1905 
Genus Schizaster L. Agassiz, 1836 

Description: Test high, sloping anteriorly from posterior vertex, beaked 
over periproct; ambulacra sunken, frontal one deeply depressed; posterior petals 
0.3 to 0.5 as long as anterior pair; apical system ethmolytic with two to four 
gonopores. 

Florida species; Two described species are reported from Florida, 
including S. armiqer (Clark, 1915) and S. ocalanus Cooke, 1942, and one 
Schizaster sp. is yet to be formally described. 

Comments; All three taxa of Schizaster are present in the Upper Ocala 


Limestone. 


Schizaster armiqer (Clark, 1915) 
(Figure 3-7, E-F) 


53 


Material examined: UF 3302 (figured test), UF 4942 (test), UF 5558 (14 
tests), UF 45568 (3 tests). 

Description: Test moderately large, much depressed and cordiform in 
marginal outline. Upper surface slopes at first rapidly from a sharp anterior 
margin to near the apical system, where it becomes nearly flat for a short 
distance. Beyond the apical system a sharp elevated ridge highest near the 
middle point continuous on to the truncated posterior margin. Ambulacra are 
broad, the single anterior ambulacrum being situated in a deep broad groove that 
deeply indents the anterior margin. Paired ambulacra have broad deep petals, 
the anterolateral being somewhat over one and a halftimes as long as the 
posterolateral. Interambulacra are more or less flat, slightly gibbous, the 
posterior much elevated forming a sharp ridge. The surface is thickly covered 
with small perforate tubercles. The peripetalous and lateral fascicles are very 
distinct. The peristome is near the anterior margin in a shallow depression. The 
periproct is high on truncated posterior margin. 

Schizaster ocalanus Cooke. 1942 
(Figure 3-7, G-H) 

Material examined: UF 5833 (figured test), UF 5864 (test), UF 41268 
(test), UF 45567 (test), UF 64439 (test). 

Description: Test subglobular, cordate, the anterior depression 
extending from the apical system to the peristome, the upper surface more 


54 


inflated than the lower. Apical system nearly central, with two large genital 
pores, one between the ends of each lateral pair of petals, and the madreporite 
extending behind them. Anterior ambulacral area moderately sunken; pores of 
each pair separated by a high granule. Petals nearly straight, sunken; anterior 
pair diverging at an angle of approximately 120°, the posterior at an angle of 
approximately 60°; anterior pair about twice as long as posterior; open at the 
distal ends; poriferous zones about as wide as interporiferous zones; pores 
conjugate. Peripetalous fascicle concave between the lateral petals, convex 
elsewhere. Peristome far forward, subtrigonal to subpentagonal, strongly lipped 
posteriorly, weakly lobate anterolaterally. Periproct about as large as the 
peristome, elliptical, higher than wide, high up on the flattened, sloping posterior 
end. Surface covered with small tubercles. 

Schizaster sp. 

(Figure 3-7, l-K) 

Material examined: UF 68927 (figured test), UF 68926 (figured test). 

Formation: Upper Ocala Limestone. 

Locality: Steinhatchee Pits (DI005); Dixie County, FL; Clara Quadrangle, 
Sec. 15/21/22/28, T8S, R10E; Hunter bed 3. 

Collector: M. Hunter. 

Date: Unknown. 

Description: Test small to medium size; horizontal outline subcircular to 
subpentagonal. Aboral surface inflated, broadly curving; highest point in 
posterior third of test along medial ridge traversing from apical system to 


55 


posterior margin. Apical system located near test center; petaloid ambulacra 
depressed, with ambulacrum III producing modest sulcus at anterior margin; 
anterior petals (II and IV) diverge at slightly less than 90° from each other as do 
posterior pair (I and V); anterior petals relatively straight, longer than posterior 
petals; terminate approximately half the distance to test margin; posterior petals 
short, extending less than half of distance to margin. Test margin broadly 
rounded at ambitus, but abrupt and flattened at posterior margin. Periproct 
small, elliptical with long axis vertical; located inframarginal just below aboral 
surface. Adoral surface slightly convex; peristome located in anterior third, small, 
transversely elliptical with modest labrum. Tubercles larger near margin, smaller 
adapically; fascicles distinct. 

Comments; Two described species, S. ocalanus and S. armiqer . are 
known from the Upper Ocala Limestone and are available for comparison with 
this fossil. The specimen described herein appears to differ somewhat from both 
of these taxa and may represent a new species. Unfortunately, the described 
fossil is not perfectly preserved, particularly in the apical system and much of the 
aboral surface, thereby limiting complete description and analysis of the 
morphological features. Recognizing these limitations, several comments are 
provided regarding possible specific identification based on the unidentified 
specimen. The peristome of S. ocalanus is very near the anterior margin, and 
proportionally closer than observed in the Schizaster sp. fossil. The unidentified 
specimen also is more strongly carinate in posterior of aboral surface as 
compared with S. ocalanus . 


56 


Comparison with S. armiqer shows differences between the fossils as 
well. The unidentified Schizaster sp. has less depressed petaloid ambulacra 
than S. armiqer , and the relative length of the petals is shorter than S. armiqer . 
Therefore, I interpret this fossil to be a new and undescribed species of 
Schizaster from the Eocene, and count this as a new taxonomic record, a unique 
species for the Eocene diversity count, and as a new stratigraphic record for 
Florida. A caution should be made, however, that the specimen is not perfectly 
preserved and more specimens must be collected to thoroughly differentiate and 
define the morphological features present on this echinoid fossil. 

Genus Aqassizia L. Agassiz and Desor, 1847 

Description: Egg-shaped, with ethmolytic apical system showing four 
gonopores and fused genital plates; frontal ambulacrum flush, petals slightly 
sunken and curiously modified; in anterior petals anterior plate row bearing tiny 
tube feet which emerge through microscopic pores, whereas pores of posterior 
plate row are normally developed; posterior petals much shorter and may be 
normal or similarly modified. 

Florida species; Two species are reported from Florida. One species is 
A. clevei Cotteau, 1875, and the second is ^ sp. cf. A. wilminqtonica Cooke, 
1942. 

Comments: A. clevei is present in the Lower Ocala Limestone, while A. 
sp. cf. A. wilminqtonica is present in the Upper Ocala Limestone. 


Aqassizia clevei Cotteau. 1875 
(Figure 3-7, L-M) 

Material examined. UF 5841 (figured test), UF 38221 (118 tests), UF 


57 


39091 (3 tests), UF 45918 (test). 

Description: Horizontal outline obovate, truncated behind; upper surface 
strongly inflated, highest in front of the genital system; margin broadly rounded; 
lower surface gently convex. Apical system behind the center; four genital pores, 
close together; ethmolytic, madreporite extending between the ocular plates. 
Anterior ambulacrum very slightly depressed. Paired petals straight, slightly 
depressed; anterior pair twice as long as the posterior; both pairs diverging at an 
angle of approximately 90°; pores of posterior petals and posterior zone of 
anterior paired petals small, pores very minute; interporiferous zones narrow. 
Peristome at the anterior third, large, reniform, weakly lipped. Periproct 
transversely oval, about midway up the posterior end, at the top of a truncation. 
Marginal fasciole curving downward below the periproct; hemipetalous fascicle 
meeting the marginal fasciole behind the anterior paired petals. 

Aqassizia sp. cf. A. wilminqtonica Cooke, 1942 
(Figure 3-7, N-0) 

Material examined; UF 68931 (figured; one incomplete test). 

Formation: Upper Ocala Limestone. 

Locality: Steinhatchee Pits (DI005); Dixie County, FL; Clara Quadrangle, 
Sec. 15/21/22/28, T8S, R10E. 


Collector; M. Hunter. 


58 


Date; Unknown. 

Description: Horizontal outline suboval to subcircular; aboral surface 
inflated, becoming less rounded in apical region; test margin broadly curved; 
adoral surface slightly convex, with longitudinal crest along plastron to posterior 
peristome labrum. Apical system positioned in slight depression, slightly anterior 
of test center; four genital pores. Anterior petals (II and IV) diverge at approxi- 
mately 105°, while posterior pair (I and V) diverge at approximately 80°; petals 
shallowly depressed, with ambulacrum III least developed and sunken; pore pairs 
not visible on specimen, but general petal shape relatively narrow and elongate. 
Periproct and posterior margin broken and absent. Peristome arcuate, about two 
to three times as wide as long; located in anterior third of test; elevated and 
prominent posterior labrum. Tubercles small, densely distributed throughout test 
surfaces. 

Comments: This fossil is an important addition to the Eocene echinoid 
record of the state. Although the specimen has undergone recrystallization and 
fragmentation, it is easy to identify the genus as Aqassizia . Comparison with 
described species within this genus tends to support my interpretation that 
matches most closely (though not perfectly) with A. wilminqtonica Cooke, 1942. 
This interpretation is supported by similar petaloid ambulacra dimensions and 
orientations, as well as the peristome shape and position found in the above 
specimen and A. wilminqtonica . However, since the newly reported fossil from 
Dixie County is incomplete, I have chosen to refer to the specimen as Aqassizia 
sp. cf. A. wilminqtonica until additional specimens are collected that show all the 


59 


morphological features necessary to confirm this interpretation. With respect to 
biostratigraphy data, the fossil is a new stratigraphic record for the Upper Ocala 
Limestone and is counted as a unique species for the Eocene diversity total, but 
it is not considered to be a new taxonomic record. 

Family Brissidae Gray, 1855 
Genus Brissiosis L. Agassiz in Agassiz and Desor, 1847 

Description: Ovate, somewhat depressed, with slight frontal sinus; 
ethmolytic, two to four gonopores; ambulacra slightly depressed; paired ones 
petaloid, may have rudimentary pores in proximal plates; petals confluent in 
some species; subanal fascicle may be lost in adults. 

Florida species: Two species present, including B. steinhatchee Cooke, 
1942 and a second, undescribed taxon referred to as Brissopsis sp. reported 
herein. 

Comments: Both species are present in the Upper Ocala Limestone. 

Brissopsis steinhatchee Cooke, 1942 
(Figure 3-7, P-Q) 

Material examined: UF 2144 (figured test), UF 4939 (4 tests), UF 4945 
(5 tests), UF 48127 (2 tests), UF 48501 (test). 

Description: Horizontal outline suboval, emarginate in front, truncate 
behind; upper surface moderately flat, sloping gently forward, with depressed 
petals and anterior ambulacrum; margin broadly rounded; lower surface gently 
convex, somewhat rostrate behind. Apical system near the anterior third; 


60 


ethmolytic; four genital pores. Petals sunken, extending about halfway to the 
margin; anterior pair straight, diverging at an angle of approximately 130°; 
posterior pair longer, curving outward to an angle of approximately 40°; both 
sunken in a single depression; pores elongate-oval; interporiferous zones nearly 
as wide as the poriferous; inner poriferous zones narrowing and becoming 
obsolete near apex. Peristome curved, with rounded ends and a posterior lip. 
Periproct large, vertical, higher than wide. 

Brissopsis sp. 

(Figure 3-8, A-B) 

Material examined: UF 68935 (test figured). 

Formation; Upper Ocala Limestone. 

Locality; Crystal River Rock Company A (CI016); Citrus County, FL; 
Flomosassa Quadrangle, Sec. 6, T19S, R18E. 

Collector: M. Hunter. 

Date: Unknown. 

Description; Horizontal outline elongate, ellipsoid; upper surface 
moderately inflated; test margin broadly rounded; adoral surface slightly convex, 
with minimally carinated posterior region. Posterior margin truncated; periproct 
located inframarginally on truncation (covered with matrix). Test partially covered 
with secondary calcite crystals and cemented matrix. Ambulacra sunken; pore 
pairs not readily visible; anterior petals longer than posterior petals, extending to 
near margin; posterior petal extending approximately half the distance to margin; 
petals relative straight. Apical system (covered, not directly visible) estimated to 


61 

lie slightly anterior of test center. Periproct and peristome covered as well. 
Tubercles generally small, with larger ones located on plastron. 

Comments: This fossil is somewhat difficult to identify with confidence as 
a result of the matrix strongly cemented to the test surface and the 
recrystallization of test plates. The only species of Brissopsis from the Eocene of 
Florida is B. steinhatchee (see earlier description). This fossil does not match 
the characteristics of B. steinhatchee in two important morphological features. 
First, the ambulacral petal length of this newly reported fossil posses anterior 
petals that are significantly longer than the posterior petals, whereas the opposite 
is true for B. steinhatchee . The second difference is in the relative position of the 
apical system, such that B. steinhatchee has its apical system in the anterior third 
of the test whereas in this fossil it is located only slightly anterior of center. Due 
to these differences, I have chosen to refer to this fossil as Brissopsis sp. herein 
and consider it to be a potential new taxonomic record, a new stratigraphic 
record for the Upper Ocala Limestone, and I include it as a unique species for the 
Eocene echinoid diversity total. 

Genus Eupataqus L. Agassiz, 1847 
Description; Test ovoid in outline, low, oral side flat; apical system 
anterior, ethmolytic, with four gonopores; paired ambulacra with closed petals; 
frontal ambulacrum non-petaloid, pores in single series, phyllodes weak; primary 
tubercles on aboral side only within peripetalous fasciole. 


62 


Florida species; Four species of Eupataqus are present, including E. 
antillarum (Cotteau, 1875), E. clevei (Cotteau, 1875), E. ocalanus Cooke, 1942, 
and one species tentatively identified as Eupataqus cf. ocalanus . 

Comments: Two species, E. antillarum and E. clevei , are present in the 
Lower Ocala Limestone and one or two species (E. ocalanus and Eupataqus sp. 
cf. E. ocalanus ) are found in the Upper Ocala Limestone. 

Eupataqus antillarum (Cotteau, 1875) 

(Figure 3-8, C-D) 

Material examined: UF 3913 (figured test), UF 46462 (test), UF 49989 
(test), UF 55191 (5 tests). 

Description: Florizontal outline obovate, truncated behind; upper surface 
moderately elevated, highest midway between the apical system and the 
periproct, sloping gently forward; lower surface flat; margin rounded. Apical 
system slightly eccentric forward, ethmolytic, the madreporite protruding far 
behind the ocular plates; four genital pores close together. Anterior ambulacrum 
narrow, not petaloid, slightly flattened, plates nearly equilateral. Petals long, 
extending nearly to the margin, slightly flexuous; anterior pair diverging at an 
angle of approximately 145°, posterior pair 40°; pores circular, strongly 
conjugate; poriferous zones nearly closed, narrow. Peristome at the anterior 
third, rounded somewhat wider than long; labrum not conspicuously projecting. 
Floscelle conspicuous. All interambulacra reaching the peristome. Periproct 
terminal, vertical, higher than wide, about mid-height, at the top of a small 
truncation. Peripetalous fasciole not indented; subanal fasciole heart shaped. 


63 


Large tubercles arranged in parallel zigzag rows in the four lateral 
interambulacra, confined within the peripetalous fasciole. Smaller tubercles 
closely cover the interambulacra on the lower side. Tubercles arranged in 
radiating rows on the escutcheon, forming a pattern suggestive of a pair of 
spread wings. Lower parts of ambulacra smooth and appearing callused. 

Eupataqus clevei (Cotteau, 1 875) 

(Figure 3-8, E-F and Figure 3-9, A-B) 

Material examined: UF 12681 (figured test), UF 12903 (figured test), UF 
35808 (test), UF 35809 (test), UF 35824 (test), UF 35825 (test). 

Description: Test large, hemispherical; horizontal outline oval; upper 
surface strongly inflated, slightly flattened in front; lower surface nearly flat; 
margin very broadly rounded. Apical system slightly anterior, small; four genital 
pores; ethmolytic, the madreporite protruding far behind the ocular plates. 
Anterior ambulacrum narrow, plates nearly equilateral, pore pairs very small. 
Petals long, extending to the margin, very wide, flush, lanceolate, ore or less 
open distally; pores oval, conjugate; poriferous zones narrow, much narrower 
than the interporiferous zones; paired ambulacra much expanded at the ends of 
the petals. Peristome at the anterior third, reniform, twice as wide as long, 
strongly lipped; floscelle conspicuous. Plates of the trivium almost excluding the 
interambulacral plates at the peristome. Periproct terminal, vertical, higher than 
wide. Peripetalous fasciole narrow, not indented. Subanal fasciole apparently 
heart shaped. Tubercles rather small, scrobiculate. 


64 

Eupataqus ocalanus Cooke, 1 942 
(Figure 3-9, C-D) 

Material examined: UF 46942 (figured test), UF 3335 (test), UF 4364 (7 
tests), UF 47043 (6 tests), UF 48489 (6 tests). 

Description; Test large, oval, truncated behind; upper surface rostrate 
above the periproct, sloping to the rounded margin; lower surface nearly flat but 
projecting downward where the subanal fasciole crosses the median line. Apical 
system slightly in front of the center; four genital pores, close together; 
madreporite extending behind the posterior pair of ocular pores. Anterior 
ambulacral area not at all petaloid, flattened. Petals slightly depressed; anterior 
pair extending four-fifths the way to the margin, diverging at an angle of 135°; 
posterior pair longer, extending three-fourths the way to the margin, diverging at 
an angle of 47°. Poriferous zones about as wide as interporiferous zones; pores 
elliptical, conjugate. Peristome large, elliptical, labiate behind, slightly farther 
forward than the apical system. Periproct large, terminal, supramarginal, pear- 
shaped. Peripetalous fasciole without indentations but having a V-shaped 
projection above the periproct. Subanal fasciole heart-shaped, enclosing a 
spread-wings-shaped escutcheon studded with large tubercles arranged in 
transverse lines. Sternum covered with somewhat smaller tubercles. Posterior 
ambulacral areas almost smooth on lower surface. Paired interambulacral areas 
on lower surface covered with a coarse reticulation of large tubercles. Large 
tubercles on upper surface confined within the peripetalous fasciole, arranged in 
zigzag rows in the posterior paired interambulacral areas, fewer and more 
scattered in the other interambulacral areas. 


65 

Eupataqus sp. cf. E. ocalanus Cooke, 1942 
(Figure 3-10, A-B) 

Material examined: UF 68936 (figured test). 

Formation: Upper Ocala Limestone. 

Locality: Crystal River Rock Company A (CI016); Citrus County, FL; 
Flomosassa Quadrangle, Sec. 6, T19S, RISE. 

Collector: M. Flunter. 

Date: Unknown. 

Description: Florizontal outline suboval, narrowing posteriorly while 
broadly curving along anterior margin. Aboral surface moderately inflated; 
highest point anterior of apical system; margins broadly curving; adoral surface 
nearly flat to slightly convex. Apical system eccentric anteriorly (not visible on 
specimen); ambulacra straight, with consistent width and rounded terminus. 
Periproct absent due to compaction and crushing. Peristome moderate in size; 
slightly wider than long; anterior edge curved, posterior edge broken; located 
approximately midway between center and anterior margin; posterior labrum 
apparently elevated and prominent. Tubercles not visible due to recrystallization. 

Comments: The specimen is in a relatively poor preservation state, 
thereby limiting the description and identification. The specimen can be identified 
as the genus Eupataqus . but the species cannot be determined at this time. A 
comparison with the species E. antillarum (Lower Ocala Limestone) does not 
match with respect to general test shape, and the same is true for E. clevei 
(Lower Ocala Limestone), which is also a significantly larger echinoid. One other 
species, E. ocalanus . is present in the Upper Ocala Limestone. The fossil 


66 


described here matches at least moderately well with E. ocalanus based on 
general test shape and peristome position. Therefore, I refer to this fossil as 
Eupataqus sp. cf. E. ocalanus to convey the uncertainty associated with the poor 
preservation. The specimen is not considered a new stratigraphic record or a 
new taxonomic record, and using the assumption that it may be another fossil of 
E. ocalanus . it is not counted as a unique species for the Eocene echinoid 
diversity total. 


Genus Macropneustes L. Agassiz, 1847 

Description: Differs from Eupataqus chiefly in having depressed petals, 
and broad test; frontal sinus distinct. 

Florida species: M- mortoni (Conrad, 1850). 

Comments: This species is present in the Upper Ocala Limestone. 

Macropneustes mortoni (Conrad. 1850) 

(Figure 3-10, C-D) 

Material examined: UF 3309 (figured test), UF 12976 (test), UF 12986 
(test), UF 38000 (test). 

Description: Test thin. Horizontal outline broadly oval, flattened or very 
slightly emarginate in front. Upper surface swollen, evenly rounded. Margin 
broadly rounded. Lower surface flatfish, depressed around the peristome. 

Apical system at the anterior third; four genital pores, close together; ethmolytic, 
the madreporite extending far behind the ocular plates. Anterior ambulacrum not 
petaloid, slightly depressed, plainly depressed near the peristome. Petals flush 


67 


or slightly depressed, straight, long, extending nearly to the margin, open distally; 
inner pores circular, outer pores oval, pores conjugate; interporiferous zones 
equal in width to the poriferous zones, not expanding medially. Peristome lunate, 
with a prominent posterior lip; at the anterior third. Periproct large, broadly oval, 
erect, higher than wide, about mid-height, at the top of a slight depression. 
Peripetalous fasciole narrow, circular in front, straight or offset behind, zigzag on 
the sides. Subanal fasciole broadly U-shaped. Large tubercles crenulate, 
perforated, not confined within the fascioles. 

Genus Plaqiobrissus Pomel, 1883 

Description; Differs from Eupataqus chiefly in having anal branches on 
subanal fasciole, long plastron, short labrum, and long, narrow, flexed petals. 

Florida species: Two species are present, including P. curvus (Cooke, 

1 942) and Plaqiobrissus ? dixie (Cooke, 1 942). 

Comments: P. curvus occurs in both the lower and upper members of 
the Ocala Limestone. Fossils of Plaqiobrissus ? dixie are present only in the 
Upper Ocala Limestone. Cooke (1959, p.87) noted the question regarding the 
true generic status of this species is a reflection of both morphology and 
preservation. He questioned the referral of this fossil to Plaqiobrissus because it 
lacks an anal fasciole (which is diagnostic of the genus), yet agreed with its 
removal from Eupataqus due to the narrowness of the ambulacra and the length 
of the plastron. Further complication of this question results from the poor 
preservation of the holotype, which is an internal mold. 


Plaqiobrissus curvus (Cooke, 1 942) 
(Figure 3-11, A) 


68 


Material examined: UF 17243 (figured test), UF 4365 (5 tests), UF 4889 
(test), UF 4966 (test), UF 48496 (test). 

Description; Test ovate, low, margin rounded. Anterior ambulacral area 
not at all petaloid, slightly depressed. Anterior pair of petals nearly as long as the 
posterior pair, widely diverging, extending more than two-thirds the way to the 
margin, constricted near the tips; interporiferous zones as wide as poriferous 
zones at the tips, twice as wide elsewhere; poriferous zones convex near the 
apex, straighter near the tip. Posterior petals curved slightly backward, diverging 
at an angle of 30°, extending less than two-thirds the way to the margin; outer 
poriferous zones convex, inner nearly straight; interporiferous zones little wider 
than poriferous zones in medial part, equal near the ends. Pores conjugate. 
Peristome oval, labiate behind, farther forward than the apical system. Periproct 
terminal, above the margin, pear-shaped. Peripetalous fasciole without lateral 
sinuations, slightly bent backward at the anterior end and more strongly bent 
backward at the posterior end. Subanal fasciole heart-shaped, enclosing a 
spread-wings-shaped escutcheon, which is studded with large tubercles arran- 
ged in transverse lines and perforated by at least six pores on each side. All four 
paired interambulacral areas studded with large tubercles within the peripetalous 
fasciole; nearly bare elsewhere on upper surface; studded with large, evenly 
spaced tubercles on lower surface. Plastron resembling fish scales. 


Plaqiobrissus ? dixie (Cooke, 1942) 
(Figure 3-11, B-C) 


69 


Material examined: UF 4929 (figured test), UF 4937 (7 tests), UF 4958 
(test), UF 4969 (test), UF 5808 (2 tests). 

Description: Test oval, depressed, truncate behind, margin rounded, 
upper surface little more inflated than the lower, rostrate above the periproct. 
Apical system in front of the center; four genital pores close together, the 
posterior pair separated by the elongated madreporite. Anterior ambulacral area 
not at all petaloid nor depressed. Petals somewhat sunken; anterior pair curved 
slightly forward, diverging at an angle of about 145°, extending about three- 
fourths of the way to the margin, shorter than the posterior pair; posterior petals 
straight, diverging at an angle of nearly 50°, extending nearly two-thirds of the 
way to the margin; poriferous zones wider than interporiferous zones; pores 
conjugate. Peristome large, wider than long, evenly curved in front, labiate 
behind. Periproct about as large as the peristome, pear-shaped, above the 
margin, on the steeply truncated end. Peripetalous fascicle without marked 
indentations. Subanal fascicle, heart-shaped, enclosing a spread-wings-shaped 
escutcheon covered with large tubercles; remainder of lower posterior 
interambulacral area covered by closer, smaller tubercles. Ambulacral areas on 
lower surface bare. Paired interambulacral areas on lower surface covered by 
lines of large tubercles. Upper surface well covered by small tubercles; large 
tubercles confined within the peripetalous fascicle, most abundant between and 
adjacent to the paired petals. 


Figure 3-1 . Eocene regular and irregular echinoids. 

A) Phvllacanthus mortoni (Conrad, 1 850); UF 66913; aboral view of test; Ocala 
Limestone; lx. 

B) Phvllacanthus mortoni (Conrad. 1850); UF 66913; adoral view of test; Ocala 
Limestone; lx. 

C) Phvllacanthus mortoni (Conrad, 1 850); UF 66913; lateral view of test; Ocala 
Limestone; lx. 

D) DIADEMATOIDA; UF 32929; hemi-pyramid of lantern; Ocala Limestone; lx. 

E) DIADEMATOIDA; UF 32929; hemi-pyramid of lantern; Ocala Limestone; lx. 

F) Dixieus dixie (Cooke, 1941); UF 5467; aboral view of test; Ocala Limestone; 
lx. 

G) Dixieus dixie (Cooke. 1941); UF 5467; adoral view of test; Ocala Limestone; 
lx. 

FI) Dixieus dixie (Cooke, 1941); UF 66559; aboral view of partial test with intact 
lantern; Ocala Limestone; lx. 

I) Olioopygus phelani Kier, 1967; UF 18017; aboral view of test; Ocala 

Limestone; lx. 

J) Oliaopyqus phelani Kier. 1967; UF 18017; adoral view of test; Ocala 
Limestone; lx. 

K) Oliaopyqus haldemani (Conrad. 1850); UF 47257; aboral view of test; Ocala 
Limestone; lx. 

L) Oliqopyqus haldemani (Conrad. 1850); UF 47257; adoral view of test; Ocala 
Limestone; lx. 

M) Oliqopyqus wetherbvi de Loriol, 1887; UF 17756; aboral view of test; Ocala 
Limestone; lx. 

N) Oliqopyqus wetherbvi de Loriol, 1 887; UF 1 7756; adoral view of test; Ocala 
Limestone; lx. 


71 



Figure 3-2. Eocene irregular echinoids. 

A) Amblvpyqus americanus Michelin, 1 858; UF 67090; aboral view of test; Ocala 
Limestone; 1x. 

B) Amblvpyqus americanus Michelin, 1858; UF 67090; adoral view of test; Ocala 
Limestone; lx. 


73 



Figure 3-3. Eocene irregular echinoids. 

A) Fibularia vauahani (Twitchell, 1 91 5); UF 38211; aboral view of test; Ocala 
Limestone; 2x. 

B) Fibularia vauahani (Twitchell. 1915); UF 38211; adoral view of test; Ocala 
Limestone; 2x. 

C) Neolaganum dalli (Twitchell. 1915); UF 104422; aboral view of test; Avon 
Park Formation; lx. 

D) Neolaganum dalli (Twitchell, 1915); UF 104422; adoral view of test; Avon 
Park Formation; lx. 

E) Neolaganum durhami Cooke, 1959; UF 13026; aboral view of test; Ocala 
Limestone; lx. 

F) Neolaganum durhami Cooke, 1 959; UF 1 3026; adoral view of test; Ocala 
Limestone; lx. 

G) Durhamella floridana (Twitchell, 1915); UF 3356; aboral view of test; Ocala 
Limestone; lx. 

FI) Durhamella floridana (Twitchell. 1915); UF 3356; adoral view of test; Ocala 
Limestone; lx. 

I) Durhamella ocalana (Cooke, 1 942); UF 3341 ; aboral view of test; Ocala 
Limestone; lx. 

J) Durhamella ocalana (Cooke. 1942); UF 3341; adoral view of test; Ocala 
Limestone; lx. 

K) Weisbordella cubae (Weisbord, 1934); UF 5846; aboral view of test; Ocala 
Limestone; lx. 

L) Weisbordella cubae (Weisbord. 1934); UF 5846; adoral view of test; Ocala 
Limestone; lx. 

M) Weisbordella iohnsoni (Twitchell, 1915); UF 47957; aboral view of test; Ocala 
Limestone; lx. 

N) Weisbordella iohnsoni (Twitchell, 1915); UF 47957; adoral view of test; Ocala 
Limestone; lx. 

O) Wvthella eldridgei (Twitchell. 1915); UF 5803; aboral view of test; Ocala 
Limestone; lx. 

P) Wvthella eldridgei (Twitchell, 1915); UF 5803; adoral view of test; Ocala 
Limestone; lx. 


75 



Figure 3-4. Eocene irregular echinoids. 

A) Periarchus Ivelli floridanus Fischer, 1951; UF 17913; aboral view of 
dolomitized, partial internal mold of test; Ocala Limestone; lx. 

B) Periarchus Ivelli floridanus Fischer, 1951 ; UF 12795; adoral view of 
dolomitized, partial internal mold of test; Ocala Limestone; lx. 


77 



Figure 3-5. Eocene irregular echinoids. 

A) Mortonella guinguefaria (Say. 1825); UF 2202a; aboral view of test; Ocala 
Limestone; lx. 

B) Mortonella guinguefaria (Say, 1825); UF 2202b; adoral view of test; Ocala 
Limestone; lx. 

C) Mortonella guinguefaria kewi (Cooke, 1942); UF 5275; aboral view of test; 
Ocala Limestone; lx. 

D) Mortonella guinguefaria kewi (Cooke, 1 942); UF 5275; adoral view of test; 
Ocala Limestone; lx. 

E) Echinolampas tanvpetalis Flarper and Shaak, 1974; UF 5385; aboral view of 
partial test; Ocala Limestone; lx. 

F) Echinolampas tanvpetalis Harper and Shaak, 1974; UF 5385; adoral view of 
partial test; Ocala Limestone; lx. 


79 



Figure 3-6. Eocene irregular echinoids. 

A) Rhvncholampas conradi (Conrad, 1 850); UF 39452; aboral view of test; 

Ocala Limestone; 1x. 

B) Rhvncholampas conradi (Conrad, 1850); UF 39452; adoral view of test; 

Ocala Limestone; lx. 

C) Rhvncholampas conradi (Conrad, 1850); UF 39452; lateral view of test; 

Ocala Limestone; lx. 

D) Rhvncholampas conradi Ivelli (Conrad, 1850); UF 3343; aboral view of test; 
Ocala Limestone; lx. 

E) Rhvncholampas conradi Ivelli (Conrad, 1 850); UF 3343; adoral view of test; 
Ocala Limestone; lx. 

F) Rhvncholampas conradi Ivelli (Conrad, 1850); UF 3343; lateral view of test; 
Ocala Limestone; lx. 

G) Rhvncholampas ericsoni (Fischer, 1951); UF 37661; aboral view of test; 
Ocala Limestone; lx. 

H) Rhvncholampas ericsoni (Fischer, 1951); UF 37661 ; adoral view of test; 

Ocala Limestone; lx. 

I) Rhvncholampas ericsoni (Fischer. 1951); UF 37661; lateral view of test; Ocala 
Limestone; lx. 

J) Rhvncholampas qeorqiensis (Twitchell, 1915); UF 22535; aboral view of 
dolomitized internal mold of test; Ocala Limestone; lx. 

K) Rhvncholampas qeorqiensis (Twitchell. 1915); UF 22535; adoral view of 
dolomitized internal mold of test; Ocala Limestone; lx. 

L) Rhvncholampas qeorqiensis qlobosus (Fischer, 1951); UF 32945; aboral view 
of dolomitized internal mold of test; Ocala Limestone; lx. 

M) Rhvncholampas qeorqiensis qlobosus (Fischer, 1951); UF 36435; aboral 
view of dolomitized internal mold of test; Ocala Limestone; lx. 

N) Rhvncholampas qeorqiensis qlobosus (Fischer, 1951); UF 36435; adoral 
view of dolomitized internal mold of test; Ocala Limestone; lx. 

O) Rhvncholampas qeorqiensis qlobosus (Fischer. 1951); UF 36435; lateral 
view of dolomitized internal mold of test; Ocala Limestone; lx. 

P) Rhvncholampas troianus (Cooke, 1942); UF 3747; aboral view of test; Ocala 
Limestone; lx. 

Q) Rhvncholampas troianus (Cooke, 1942); UF 3747; adoral view of test; Ocala 
Limestone; lx. 


81 



Figure 3-7. Eocene irregular echinoids. 

A) Eurhodia patelliformis (Bouve, 1 851 ); UF 4932; aboral view of test; Ocala 
Limestone; 1x. 

B) Eurhodia patelliformis (Bouve, 1851); UF 4932; adoral view of test; Ocala 
Limestone; lx. 

C) Ditremaster becked (Cooke. 1942); UF 47040; aboral view of test; Ocala 
Limestone; lx. 

D) Ditremaster beckeri (Cooke, 1942); UF 47040; adoral view of test; Ocala 
Limestone; lx. 

E) Schizaster armiqer (Clark, 1915); UF 3302; aboral view of test; Ocala 
Limestone; lx. 

F) Schizaster armiqer (Clark, 1915); UF 3302; adoral view of test; Ocala 
Limestone; lx. 

G) Schizaster ocalanus Cooke, 1942; UF 5833; aboral view of test; Ocala 
Limestone; lx. 

FI) Schizaster ocalanus Cooke. 1942; UF 5833; adoral view of test; Ocala 
Limestone; lx. 

I) Schizaster sp.; UF 68927; aboral view of test; Ocala Limestone; lx. 

J) Schizaster sp.; UF 68927; adoral view of test; Ocala Limestone; lx. 

K) Schizaster sp.; UF 68926; aboral view of test; Ocala Limestone; lx. 

L) Aaassizia clevei Cotteau, 1875; UF 5841; aboral view of test; Ocala 
Limestone; lx. 

M) Aaassizia clevei Cotteau, 1875; UF 5841; adoral view of test; Ocala 
Limestone; lx. 

N) Aaassizia sp. cf. A. wilmingtonica Cook, 1942; UF 68931; aboral view of 
partial test; Ocala Limestone; lx. 

O) Aaassizia sp. cf. A. wilmingtonica Cook, 1942; UF 68931; adoral view of 
partial test; Ocala Limestone; lx. 

P) Brissopsis steinhatchee Cooke, 1942; UF 2144; aboral view of test; Ocala 
Limestone; lx. 

01 Brissopsis steinhatchee Cooke. 1942; UF 2144; adoral view of test; Ocala 
Limestone; lx. 


83 



Figure 3-8. Eocene irregular echinoids. 

A) Brissopsis sp.; UF 68935; aboral view of partial test; Ocala Limestone; 1x. 

B) Brissopsis sp.; UF 68935; adoral view of partial test; Ocala Limestone; 1x. 

C) Eupataqus antillarum (Cotteau, 1875); UF 3913; aboral view of test; Ocala 
Limestone; lx. 

D) Eupataqus antillarum (Cotteau. 1875); UF 3913; adoral view of test; Ocala 
Limestone; lx. 

E) Eupataqus clevei (Cotteau, 1875); UF 12681; aboral view of dolomitized 
internal mold of test; Ocala Limestone; lx. 

F) Eupataqus clevei (Cotteau. 1875); UF 12681; adoral view of dolomitized 
internal mold of test; Ocala Limestone; lx. 


85 









Figure 3-9. Eocene irregular echinoids. 

A) Eupataqus clevei (Cotteau, 1875); UF 12903; aboral view of test; Ocala 
Limestone; lx. 

B) Eupataqus clevei (Cotteau, 1875); UF 12903; adoral view of test; Ocala 
Limestone; lx. 

C) Eupataqus ocalanus Cooke, 1942; UF 46942; aboral view of test; Ocala 
Limestone; lx. 

D) Eupataqus ocalanus Cooke, 1942; UF 46942; adoral view of test; Ocala 
Limestone; lx. 


87 



Figure 3-10. Eocene irregular echinoids. 

A) Eupataqus sp. cf. E. ocalanus Cooke, 1942; UF 68936; aboral view of 
limestone encrusted test; Ocala Limestone; lx. 

B) Eupataqus sp. cf. E. ocalanus Cooke, 1942; UF 68936; adoral view of 
limestone encrusted test; Ocala Limestone; lx. 

C) Macropneustes mortoni (Conrad, 1850); UF 3309; aboral view of test; Ocala 
Limestone; lx. 

D) Macropneustes mortoni (Conrad, 1 850); UF 3309; adoral view of test; Ocala 
Limestone; lx. 


89 




Figure 3-11. Eocene irregular echinoids. 

A) Plaaiobrissus curvus (Cooke. 1942); UF 17243; aboral view of dolomitized 
internal mold of test; Ocala Limestone; lx. 

B) Plaaiobrissus ? dixie (Cooke, 1942); UF 4929; aboral view of partial test; 
Ocala Limestone; lx. 

C) Plaaiobrissus ? dixie (Cooke, 1942); UF 4929; adoral view of partial test; 
Ocala Limestone; lx. 


91 



92 


Oligocene Echinoids 

Order Phymosomatoida Mortensen, 1904 
Family Stomechinidae Pomel, 1883 
Genus Phymotaxis Lambert and Thiery, 1914 

Description; Test low hemispherical, medium-sized. Ambulacra 
polyporous, pore pairs in double series adorally, in single undulating line 
adapically. Primary tubercles in two regular series in each area. 

Florida species: P. mansfieldi Cooke, 1941 and Phymotaxis sp. 

Comments: Both species are present in the Suwannee Limestone. The 
unidentified species has been examined by seyerai echinoid workers in addition 
to me, and the consensus interpretation is that it is not P. mansfieldi . The fossil 
awaits further detailed examination and description, but I haye included this in 
both the new stratigraphic and taxonomic records categories. 

Phymotaxis mansfieldi Cooke, 1941 
(Figure 3-12, A-B) 

Material examined; UF3344 (figured test), UF 3325 (test). 

Description: Test large, nearly circular; upper surface somewhat 
depressed; lower surface eyenly rounded, concaye near the peristome. 
Ambulacral areas little more than half as wide as interambulacral areas; pores on 
upper surface and on ambitus arranged in connected arcs, six to eight pairs on 
each plate; pore pairs crowded on lower surface. Tubercles large, imperforate; 
two rows on each ambulacral area, four rows on each interambulacral area at the 
ambitus, the lateral rows dwindling and disappearing away from the ambitus; 
each primary tubercle surrounded by a line of small tubercles that generally 


93 


follow the edges of the plates, being somewhat larger than the others. Peristome 
small, subdecagonal; gill slits about as deep as wide, callous. 

Phymotaxis sp. 

(Figure 3-12, C) 

Material examined; UF13047 (figured test). 

Formation; Suwannee Limestone. 

Locality; Alachua County, FL; Gainesville East Quadrangle, SE1/4, 

T10S, R19E. 

Collector; R. Portell. 

Date; 3/4/81. 

Description; Horizontal outline circular; test large, subhemispherical, with 
evenly rounded margin; low in profile. Petaloid ambulacra convergent at apical 
system and peristome; maximum width at ambitus; zygopores close, rounded; 
ambulacra approximately two-thirds as wide as interambulacra. Two rows of 
prominent spines in both ambulacral and interambulacra zones; only scrobicular 
ring showing secondary spines. Apical system not visible due to recrystallization 
and replacement. Peristome large, circular, approximately one-half test 
diameter. 

Comments; This fossil is moderately well-preserved, but degraded from 
its original state via silicification and recrystallization of the calcite as well as 
dissolution of portions of the test. Slight compaction likely also altered the 
specimen from its original form. However, it is identifiable to the generic level 
and I refer to this specimen as Phymotaxis sp. Comparison with the previously 


94 


documented species, P. mansfieldi Cooke, 1941 (see description in this section) 
reveals differences between the fossils. One such difference is the peristome 
diameter. The diameter in P. mansfieldi is relatively small in contrast with that of 
this unidentified species. The general shape of the test is taller in P. mansfieldi 
whereas Phymotaxis sp. is lower with a shield-like shape. The limited detail 
preserved in this fossil prevents a more thorough comparison with other 
described species, but based on what is observed, it is likely this fossil is a new 
species of Phymotaxis . Therefore, I include it as a new stratigraphic record for 
the Suwannee Limestone, a new taxonomic record, and include it in the total 
echinoid diversity value for the Oligocene of Florida. 

Order Temnopleuroida Mortensen, 1942 
Family uncertain 
Genus Gaqaria Duncan, 1889 

Description; Moderate-sized, low hemispherical. Angular pores and pits 
lacking. No distinct sculpture on test. Tubercles crenulate, not indented, forming 
regular series in both areas. Apical system with only ocular I insert. 

Florida species: G. mossomi (Cooke, 1941). 

Comments: The species is present in the Suwannee Limestone. Speci- 
mens consist of tests, fragments, and spines in varying states of preservation. 

Gaqaria mossomi (Cooke. 1941) 

(Figure 3-12, D-E) 

Material examined: UF 28245 (figured test), UF 32350 (3 tests), UF 


32401 (2 tests), UF 32531 (6 tests). 


95 


Description: Test rather small, subhemispherical, upper surface slightly 
flattened, lower surface rounded below the ambitus and slightly concave around 
the large notched peristome. Ambulacral areas narrower than interambulacral 
areas. Ambulacral pores large; pairs uniserial, nearly straight, but each group of 
three pores slightly curved around a large tubercle; each pore pair divided by a 
raised septum. Interporiferous areas provided with two rows, about 20 in each 
row, of moderately strong, crenulated, imperforate primary tubercles and two 
shorter rows of smaller tubercles. Interambulacra provided with two rows, about 
16 in each row, of primary tubercles about equal in size to those on the 
ambulacra and with several rows of smaller tubercles, which are most numerous 
on the ambitus. 


Order Clypeasteroida A. Agassiz, 1872 
Family Clypeasteridae L. Agassiz, 1835 
Genus CIvpeaster Lamarck, 1801 

Description; Medium-sized to large, test flattened to highly campanulate, 
margin rounded to flattened and inflated; peristome usually in deep infundibulum; 
oral surface flat to concave; petals variable, closed and rounded to open or 
sublyrate, with outer pores elongate, inner ones rounded, commonly connected 
by groove; periproct usually inframarginal, rarely marginal, situated between third 
and fourth, or fourth and fifth pair of coronal plates; buccal membrane naked, 
with imbedded irregular spicules; internal supports variable in abundance, 
consisting of thin laminae and pillars; wall of test sometimes double, separated 
by pillars. Variation in external test morphology and shape of petals is very 


96 

great, more than 400 nominal taxa existing in the literature, but no systematic 
basis for subgeneric groupings can be recognized. 

Florida species: Four species are present, including C. batheri Lambert 
1915, C. cotteaui Egozcue, 1897, C. oxybaphon Jackson, 1922, and C. roqersi 
(Morton, 1834). 

Comments; All four species are present in the Suwannee Limestone. In 
addition to their presence the Suwannee Limestone, C. cotteaui is preserved in 
the Bridgeboro Limestone while C. roqersi is found in the Marianna Limestone. 
The fossils of C. batheri reported herein represent a new stratigraphic occurrence 
of this species in the Suwannee Limestone. 

CIvpeaster batheri Lambert, 1915 
(Figure 3-12, F-G) 

Material examined: UF 2546 (figured test), UF 5341 (test). 

Description: Test medium size, longer than wide, depressed, oval, 
narrowed anteriorly, subtruncated posteriorly. Upper surface moderately convex, 
ventrally widely and deeply concave. Petals wide open, with a tendency to 
narrow toward the extremities. 

CIvpeaster cotteaui Egozcue, 1897 
(Figure 3-12, H-l) 

Material examined; UF 54993 (figured test), UF 54996 (test), UF 54997 
(test), UF 54998 (test fragment), UF 54999 (test fragment), UF 67416 (test). 


97 

Description: Test moderate size, oval, narrowed anteriorly, upper 
surface slightly convex. Margin thick and rounded. Lower surface concave, with 
ambulacral furrows which expand and deepen approaching peristome. 

Peristome pentagonal. Periproct circular, slightly separated from the margin. 
Apex nearly central, only slightly eccentric anteriorly. Apical disk small, with 
madreporite constituting a relatively large star. Ambulacral areas superficial, 
petals truncate and very open at the distal tips. Poriferous areas slightly 
depressed and very characteristic, the inner series of pores curving and 
converging near the apex of each ambulacrum, below making a straight line to 
distal tips, but converging. Outer series of pores curve throughout entire length, 
forming in the extreme tip a curve of less radius, converging to meet the inner 
series: as a sort of termination, four pores, smaller and elongated, form an 
irregular quadrilateral. Each poriferous plate with six tubercles. 

CIvpeaster oxvbaphon Jackson. 1922 
(Figure 3-13, A-B) 

Material examined: UF 4926 (figured test). 

Description: Test low, pentagonal in outline, elongate anteriorly, truncate 
posteriorly; thick and rounded on the margin, depressed in a dish-like fashion 
dorsally, and in the center of the dish the proximal portion of the petals with the 
apical disk rise in a slight eminence, moderately concave ventrally. Ambulacral 
petals slightly raised above the general surface of the test; petals gently rounded, 
but open at the distal tips. Poriferous areas are sunken; pores are connected by 
well-marked furrows, the ridges between poriferous areas present a gentle curve 


98 


from the base to the distal end. Petals I, V, and III are closely of the same length 
and width; II and IV are shorter, but of about the same width as the others. The 
apical disk is central in position, genital pores are large and sunken. Periproct 
well removed from the posterior border of the test. Ambulacral grooves are 
strongly marked, deepening toward the centrally placed mouth. Tubercles are 
small, closely associated, and somewhat larger and more crowded on the ventral 
side. 


CIvpeaster roqersi (Morton, 1834) 

(Figure 3-13, C-D) 

Material examined; UF 3314 (figured test), UF 65819 (7 tests), UF 
65820 (6 tests), UF 68932 (test). 

Description: Test outline oval to subpentagonal; upper surface low to 
tumid apically; lower surface relatively flat, slightly concave near the peristome; 
margin usually thin. Apical system central, tumid, with five gonopores. Petals 
extending more than halfway to the margin, completely closed apically, wide 
open distally; poriferous zones somewhat narrower than the interporiferous, inner 
pores circular, outer pores slightly elongated, pores weakly conjugate; inner side 
of poriferous zones almost straight, outer side slightly convex. Peristome central, 
pentagonal, sides slightly swollen, like bourrelets. Ambulacral grooves straight, 
narrow, extending to the margin. Periproct circular, located about one-fifth the 
distance from the margin to the peristome. Tubercles on upper surface small, 
sunken; tubercles on lower surface larger, in much larger scrobicules; 
intermediate spaces pitted. 


99 


Order Cassiduloida Claus, 1880 
Family Cassidulidae L. Agassiz and Desor, 1847 
Genus Rhvncholampas A. Agassiz, 1869 

Description: See Eocene echinoid section for generic description. 

Florida species: R. qouldii (Bouve, 1846). 

Comments: This species is present in the Suwannee Limestone and 
represents a relatively abundant and pervasive macrofossil in the formation. The 
biostratigraphic value of this species is strong because of the clear and 
unquestioned association with most facies of this Oligocene limestone unit. 

Rhvncholampas qouldii (Bouve, 1846) 

(Figure 3-13, E-F) 

Material examined: UF 67813 (figured test), UF 36608 (test), UF 46464 
(6 tests), UF 46632 (6 tests), UF 66562 (test). 

Description: Aboral test surface conico-convex, more sloping posteriorly 
than anteriorly. Margin somewhat rounded except near and under anus, where 
by a depression it becomes acute. Adoral surface subcircular; peristome located 
about one-third the distance from anterior margin. Apex subcentral, slightly 
anterior, though not as far as peristome. Ambulacra radiating at unequal angles, 
the interambulacral spaces dividing the three anterior from the two posterior 
being wider than the rest. Ambulacral pores in pore pairs diverge considerably 
from the apex, becoming quite dilated a short distance from apical system, then 
converging as they descend, until about two-thirds the distance from the peak to 
the margin; pores then change from double rows to single; at margin pores again 
dilate, and are traceable to termination at peristome where distinctly prominent. 


Anterior ambulacrum much narrower than rest. Periproct transverse, situated 
about one-fifth the distance from posterior margin to apex. 


100 


Order Spatangoida Claus, 1876 
Family Schizasteridae Lambert, 1905 
Genus Schizaster L. Agassiz, 1836 

Description: See Eocene echinoid section for generic description. 

Florida species; S. americanus Clark, 1915. 

Comments; This species is present in two formations (possibly three with 
the Bridgeboro Limestone), including the Suwannee Limestone and the Marianna 
Limestone. Its presence in both of the formations reported herein thereby 
reflects two new stratigraphic records for this species in Florida. 

Schizaster americanus Clark. 1915 
(Figure 3-13, G-H) 

Material examined; UF 55006 (figured test), UF 5276 (2 tests), UF 
27345 (test). 

Description: Test of medium size, rather tall, subpentagonal, as wide as 
long, sloping up from the anterior margin to the nearly central apical system, 
beyond which a sharp rise continues toward the posterior margin, its highest 
point being about midway. Ambulacra narrow; the anterior one situated in a 
deep, moderately wide groove that indents the anterior margin. Paired 
ambulacra with deep short petals, the anterolateral being about twice as long as 
the posterolateral. Interambulacra are broad and somewhat gibbous on the 
sides; posterior interambulacrum is much elevated and rather narrow. Test 


101 


surface covered with numerous small, but clearly distinct tubercles with small 
granules between them. The peripetalous and lateral fascicle can be readily 
traced. Apical system small, nearly central in position. Peristome near anterior 
margin in a well-marked depression. Periproct high on truncated posterior 
margin. 


Genus Aqassizia L. Agassiz and Desor, 1847 

Description: See Eocene echinoid section for generic description. 

Florida species: A. mossomi Cooke, 1942. 

Comments: This species is present in the Suwannee Limestone. 

Aqassizia mossomi Cooke, 1942 
(Figure 3-13, l-J) 

Material examined: UF 55007 (figured test), UF 3316 (2 tests), UF 
27143 (tests), UF 27183 (7 tests), UF 27187 (4 tests). 

Description: Test nearly spherical, less tumid beneath, truncated behind. 
Apical system nearly central, with four genital pores, the posterior pair separated 
by an elongated madreporite, which extends behind them. Petals forming a 
nearly right-angled X; more deeply sunken than customary for the genus; front 
pair half again as long as the back pair; pores of anterior zone of front pair small 
and oblique distally, very minute near the apex; interporiferous zones of back pair 
narrower than the poriferous zones. Anterior ambulacral area slightly sunken 
near the apex, flush at the margin. Peristome semilunate, with a posterior lip, 
located at the anterior third. Periproct large, transversely oval, terminal, at the 


102 

top of the truncation. Marginal fasciole forming a V-shape below the periproct, 
joined by the hemipetalous fasciole behind the anterior pair of petals. 
Interambulacral areas covered with close-set tubercles. 

Family Brissidae Gray, 1855 

Genus Brissopsis L. Agassiz in Agassiz and Desor, 1847 
Description: See Eocene echinoid section for generic description. 

Florida species: Brissopsis sp. 

Comments: This species is present in the Suwannee Limestone, 
represented by only two specimens. My tentative interpretation is that they likely 
represent a new species, but further detailed examination and collection of 
additional specimens may be required to formally describe these fossils. 

Flowever, I do include these samples in my taxonomic list and I believe they 
represent both new taxonomic and new stratigraphic records for the Suwannee 
Limestone. 

Brissopsis sp. 

(Figure 3-13, K-L) 

Material examined: UF 9195 (figured test), UF 10719 (figured test). 
Formation: Suwannee Limestone. 

Locality: Flernando Beach 01 (FIE001); Flernando County, FL; Aripeka 
Quadrangle, NE1/4, Sec. 24, T23S, R16E. 

Collectors: R. Portell (UF 9195) and J. Pendergraft (UF 10719). 

Date: 2/8/87 and 4/16/87, respectively. 

Description: Both specimens still embedded in matrix, thereby limiting 


103 


observation. Test small; aboral surface hemispherical with peak at apical 
system; small tubercles apparent but poorly preserved. Apical system small; four 
gonopores; petaloid ambulacra sunken; petals diverging, interpolated to be at 
nearly right angles. 

Comments; The small area of test exposed above matrix in both fossils 
along with recrystallization of the calcite plates and fractured regions prevents 
specific identification. The general test shape visible, sunken ambulacra, and 
four gonopores allows the tentative identification as Brissopsis sp. This 
preliminary identification is tenuous, however, since no other Brissopsis species 
have been reported from the Oligocene of Florida or neighboring states. The first 
record of Brissopsis from the Eocene of Florida also is part of this study, and thus 
comparison materials are not available in the Florida Museum of Natural History 
or other repositories. One record of Brissopsis in the Caribbean region is a 
report of an Oligocene species, B. aquavoi Sanchez Roig, 1952, from Cuba. 
Therefore, it is not unreasonable to expect the genus to be present in Florida 
strata as well. I include this fossil as a new stratigraphic record for the Suwan- 
nee Limestone, and assume the identification to valid at the generic level, 
thereby also permitting an interpretation of this fossil as a new taxonomic record. 


Figure 3-12. Oligocene regular and irregular echinoids. 

A) Phvmotaxis mansfieldi Cooke. 1941; UF 3344; lateral view of test; Suwannee 
Limestone; lx. 

B) Phvmotaxis mansfieldi Cooke, 1941; UF 3344; adoral view of test; Suwannee 
Limestone; lx. 

C) Phvmotaxis sp.; UF 13047; aboral view of silicified test; Suwannee 
Limestone; lx. 

D) Gaqaria mossomi (Cooke. 1941); UF 28245; aboral view of test; Suwannee 
Limestone; lx. 

E) Gaqaria mossomi (Cooke, 1 941 ); UF 28245; adoral view of test; Suwannee 
Limestone; lx. 

F) CIvpeaster batheri Lambert. 1915; UF 2546; aboral view of test; Suwannee 
Limestone; 0.75x. 

G) CIvpeaster batheri Lambert, 1915; UF 2546; adoral view of test; Suwannee 
Limestone; 0.75x. 

FI) CIvpeaster cotteaui Egozcue, 1897; UF 54993; aboral view of test; 

Bridgeboro Limestone; lx. 

I) CIvpeaster cotteaui Eqozcue. 1897; UF 54993; adoral view of test; Bridgeboro 
Limestone; lx. 


105 







Figure 3-13. Oligocene irregular echinoids. 

A) CIvpeaster oxybaphon Jackson, 1 922; UF 4926; aboral view of partial test; 
Suwannee Limestone; lx. 

B) CIvpeaster oxybaphon Jackson, 1922; UF 4926; adoral view of partial test; 
Suwannee Limestone; lx. 

C) CIvpeaster roqersi (Morton. 1834); UF 3314; aboral view of test; Suwannee 
Limestone; lx. 

D) CIvpeaster roqersi (Morton, 1 834); UF 3314; adoral view of test; Suwannee 
Limestone; lx. 

E) Rhvncholampas qouldii (Bouve, 1 846); UF 67813; aboral view of test; 
Suwannee Limestone; lx. 

F) Rhvncholampas qouldii (Bouve, 1 846); UF 6781 3; adoral view of test; 
Suwannee Limestone; lx. 

G) Schizaster americanus Clark, 1915; UF 55006; aboral view of test; 
Bridgeboro Limestone; lx. 

H) Schizaster americanus Clark, 1915; UF 55006; adoral view of test; 
Bridgeboro Limestone; lx. 

I) Aqassizia mossomi Cooke, 1942; UF 55007; aboral view of test; Suwannee 

Limestone; lx. 

J) Aqassizia mossomi Cooke. 1942; UF 55007; adoral view of test; Suwannee 
Limestone; lx. 

K) Brissopsis sp.; UF 9195; aboral view of test imbedded in limestone; 
Suwannee Limestone; lx. 

L) Brissopsis sp.; UF 10719; aboral view of test imbedded in limestone; 
Suwannee Limestone; lx. 


107 




108 

Miocene Echinoids 

Order Cidaroida Claus, 1880 
Family Cidaridae Gray, 1825 
Genus Prionocidaris A. Agassiz, 1 863 

Description: Test arched or low, more or less flattened at apex, thin and 
somewhat fragile. Primary tubercles noncrenulate adorally, weakly subcrenulate 
or noncrenulate aborally; areoles shallow, well separated save for lowermost two 
or three, which may be confluent. Pores distinctly conjugate or subconjugate. 
Primary spines usually long, tapering, with coarse thorns in longitudinal series; 
less commonly cylindrical, smooth or widened distally, or with thorns arranged in 
whorls; cortex thin; oral primaries with relatively long collar, tipped by rudimentary 
shaft. Secondary spines not adpressed, larger ones flattened, smaller ones 
spiniform. Tridentate pedicellariae slender; large globiferous pedicellariae 
without end tooth, or wanting; small globiferous with end tooth. 

Florida species; P. cookei Cutress, 1976. 

Comments: The species occurs in the Chipola and Torreya formations of 
the Florida panhandle. The specimens from the Torreya Formation are new 
stratigraphic records for the state, and are dominated by spines. These warrant 
closer examination to determine if they represent a new species. 

Prionocidaris cookei Cutress, 1976 
(Figure 3-14, A-C) 

Material examined; UF 66632 (figured test fragment), UF 88540 (figured 
radioles), UF 101422 (figured test plates), UF 44645 (12 interambulacral plates), 
UF 44660 (4 radioles). 


109 


Description: Test medium size. Primary areoles more than half width of 
plate, those near peristome confluent. Scrobicular tubercles round, separated; 
secondary tubercles otherwise well spaced, of decreasing size to sutures, not 
horizontally aligned. Radioles moderately long, slender, tapered, sometimes 
dark-banded, and with regular longitudinal series of uniformly small or small and 
medium sized spinules. Collar often long, mottled, with low, oval nodules. Oral 
radioles not capped. 

Gen. et sp. indet. 

(Figure 3-14, D) 

Material examined: UF 66579 (figured internal mold of single 
interambulacrum). 

Formation: Chattahoochee Formation (uncertain). 

Locality: Dry Creek 01 (JA010); Jackson County, FL; Oakdale 
Quadrangle, NE1/4, SW1/4, Sec. 11, T3N, R10W. 

Collectors: T. Cassady and B. Shumaker. 

Date: 7/10/93. 

Description: Specimen consists of a poorly preserved internal mold of a 
single interambulacrum. Test medium size (portion preserved with height of 
approximately 35 mm), likely spherical. Interambulacral plates about twice as 
wide as long; slightly inflated. Few pore pairs visible adjacent to segments of 
adradial suture; zygopores transverse, elliptical shape, small, and closely 
spaced; pores appear relatively arranged in relatively straight column. 

Comments: Overall preservation is poor, yet the general shape of the 
interambulacrum and its plates tends to indicate a unique regular echinoid. A 


110 


cidaroid species, Prionocidaris cookei . was described from the Miocene from 
Florida by Cutress (1976), and they have been reported from various Miocene 
units in the Caribbean (Cutress, 1980; Donovan, 1993). In Florida, cidaroid 
species are present in the Eocene ( Phyllacanthus mortoni . Ocala Limestone), 
Miocene ( Prionocidaris cookei , Chipola Formation), and Pliocene (Eucidaris 
tribuloides , Tamiami Formation). This fossil is too incomplete to allow a generic 
or specific identification to be proposed, but the familial assignment is reasonable 
based on the general arrangement and shape of the test plates as well as the 
pore pair orientation. Therefore, this fossil is a new stratigraphic record for the 
Chattahoochee Formation, and very likely a new taxonomic record for Florida as 
well since it is has very little similarity to the Chipola Formation species. 

Gen. et sp. indet. 

(Figure 3-14, E) 

Material examined. -UF 66513 (figured; one incomplete radiole). 

Formation: Shoal River Formation. 

Locality: Shoal River Grotto (WL004); Walton County, FL; New Flarmony 
Quadrangle, Sec. 4, T3N, R21W. 

Collectors: G. Schmelz and W. Conway. 

Date: 4-5/5/93 

Description: The fossil consists of a single, incomplete radiole. Both 
proximal and distal ends missing (including base, collar, and milled ring). 
Longitudinal profile straight, with only very slight narrowing toward distal end. 
Shaft ornamented with multiple longitudinal rows of very small spinelets or 


111 


thorns, all oriented toward distal end; spinelets estimated at 0.1 to 0.2 mm long. 
Overall length of radiole is 18 mm. 

Comments: identification of the lower taxonomic levels for this fossil is 
not possible based only on this radiole. The spine has morphological 
characteristics such as the rows of spinelets that reflect features common to the 
cidaroids, and therefore I believe it is reasonable to assume this ordinal 
affiliation. However, without a more complete spine or additional tests or test 
fragments, more detailed identification cannot be completed. This fossil is 
included as a new stratigraphic record for the Shoal River Formation and as one 
of the taxa from the Miocene (but not as a new taxonomic record). 

Order Arbacioida Gregory, 1900 
Family Arbaciidae Gray, 1855 
Genus Arbia Cooke, 1948 

Description; Apical system probably dicyclic, pentagonal, encircling the 
periproct. Ambulacra narrower than the interambulacral areas throughout; porie- 
rous zones uniserial, the zygopores forming a straight line above the ambius, 
arranged in slightly diagonal groups of three at and below it, not at all expaned at 
the peristome; zygopores one to each plate near the apex, three to each 
compound plate below; without sphaeridial pits. Tubercles imperforate, more 
abundant below the ambitus. Peristome small, circular, moderately notched. 


Florida species: Arbia sp. 


112 


Comments; One unidentified species, herein referred to as Arbia sp., is 
present in the Chattahoochee Formation. This taxon is new to the Florida fossil 
record. 

Arbia sp. 

(Figure 3-14, F-G) 

Material examined: UF 102309 (figured partial external mold (with 
associated RTV silicone rubber peel generated from the mold). 

Formation; Chattahoochee Formation; beds 6 and 7. 

Locality: Jim Woodruff Dam (JA003); Jackson County, FL; 
Chattahoochee Quadrangle, N1/2, Sec. 31, T4N, ROW. 

Collectors: R. Portell, C. Oyen, and L. Anderson. 

Date: 6/5/97. 

Description: Test small to moderate size (mold corona from lower 
ambitus to near apical system approximately 17 mm tall); aboral surface 
subhemispherical in shape. Two interambulacral rows of primary tubercles; 
prominent boss and mamelon (possibly perforate); distinct scrobicular ring; two 
ambulacral rows of moderate marginal tubercles; secondary tubercles limited to 
area above and below ambitus. Interambulacral plates devoid of secondary 
ornamentation nearing apical system. Compound ambulacral plates with five to 
six zygopores each; pore pairs small, circular, and closely spaced, producing 
straight columns. No remnant of apical system, periproct, peristome, or aboral 
surface is present in the specimen. 

Comments: This fossil consists only of an external mold of a portion of 
the aboral surface, therefore detailed analysis of its features is somewhat limited. 


113 


It should be noted, however, that the silicone rubber peel produced from the mold 
does indeed show the micro-morphology of that portion of the test that produced 
the mold. Based on close examination of features on both the mold and the peel, 
two different genera are possible matches for the fossil’s characteristics. One 
potential match is Gaqaria and the second is Arbia . A problem with using both of 
these genera as matches is that neither have been reported from the Miocene of 
Florida (although I also have described the affinity of a second unidentified 
regular echinoid fragment and spines from the Miocene Parachucia Formation 
with Gaqaria in this dissertation). Cooke (1948) erected the genus Arbia based 
on fossils from the late Oligocene Chickasawhay Limestone in Alabama and the 
early Miocene Paynes Hammock Formation of Mississippi (Cooke, 1959). Since 
the Florida fossil described above is Miocene and matches some of the 
characteristics (at least those which are preserved in the moldic fossil) of Arbia . I 
have chosen to refer to the fossil as a questionable Arbia sp. until additional 
fossils can be collected and used for comparison. 

Order Temnopleuroida Mortensen, 1942 
Family uncertain 
Genus Gaqaria Duncan, 1889 

Description: See Oligocene echinoid section for generic description. 

Florida species; As many as three, undescribed species herein referred 
to as Gaqaria sp. 

Comments: These species are present in the Chattahoochee and 


Parachucia formations. 


114 

Gaqaria sp. or Gaqaria spp. 

(Figure 3-14, H) 

Material examined: UF 25339 (figured test fragment), UF 25130 (90 
radioles or radiole fragments), UF 25498 (50 test fragments), UF 25514 (150 
radioles). 

Formation: Parachucia Formation. 

Locality: White Springs (HA001); Flamilton/Columbia County, FL; White 
Springs West Quadrangle; W1/2, NW1/4, SW1/4, Sec. 7, T2S, R16E for UF 
25339 and NW1/4, NW1/4, SW1/4, Sec. 7, T2S, R16E for UF 25130. 

Collectors: R. Portell and G. Morgan (UF 25339), and G. Morgan (UF 

25130) 

Date: 1/13/89 (UF 25339) and 2/13/88 (UF 25130). 

Description: The test fragment consists of a single, incomplete 
ambulacrum plate series, but is well-preserved and exhibits micro-morphological 
features clearly. Interporiferous zone with moderate-sized, crenulated marginal 
tubercles; two rows of small, non-ornate, inner tubercles adjacent to perradial 
suture; marginal tubercles scrobiculate with pronounced boss and mamelon. 

Pore pairs circular, nonconjugate with distinct septum; three or possibly four pairs 
per plate. Test fragment size limited to approximately 8x12 mm, and curvature of 
preserved section implies the overall size of complete test was small. Radioles 
small (maximum length approximately 17 mm), ranging from ovate to semi- 
bladed transversely; acetabulum conical; base smooth with narrow ring distally; 
most with smooth to slightly ornamented shaft. 


115 


Comments: Due to the limitations of identification based on this small 
and incomplete test fragment, only a tentative identification is offered as a 
potential species of Gaqaria . This is an important biostratigraphic record 
because it represents the first Miocene record of this genus in the southeastern 
U.S. Although specific assignment is unlikely based on the current fossils, it is 
probable it also is a new species as well, and thereby increases the taxonomic 
record of the Miocene in Florida too. 

Unidentified regular echinoid; cf. Gaqaria sp. 

(Figure 3-14, l-J) 

Material examined; UF 60668 (figured incomplete internal mold). 
Formation: Chattahoochee Formation; beds 6 and 7. 

Locality: Jim Woodruff Dam (JA003); Jackson County, FL; 
Chattahoochee Quadrangle, N1/2, Sec. 31, T4N, R6W. 

Collectors: R. Portell and J. Bryan 

Date: 11/18/92. 

Description: Incomplete internal mold of test; horizontal outline 
subpentagonal. Test small, subhemispherical, slightly concave adoral surface. 
Peristome approximately half to slightly greater than half of test diameter. Pore 
pairs small, round, closely set; produce ambulacra that become close together 
near peristome but diverge significantly at ambitus and higher on adapical 
surface. 

Comments: Lower taxonomic identification of this specimen is not 
possible based on the preservation of the specimen. The overall test shape, the 


116 


subpentagonal outline, and the small pore pairs may indicate an affinity to 
Gaqaria . though this can only be considered tentative at best. Clearly, the 
specimen is a regular echinoid, thereby representing the first stratigraphic record 
of such an echinoid within the Chattahoochee Formation. It also is possible this 
may be a new taxa since no Gaqaria have been reported from the Miocene 
(except in this dissertation; see earlier Gaqaria sp. description for fossils from the 
Parachucia Formation). 


Family Echinidae Gray, 1825 
Genus Psammechinus L. Agassiz and Desor, 1846 

Description: Widest at circular ambitus; ambulacral plates trigeminate, 
with primary tubercle on each; buccal membrane densely plated, with contiguous 
or even imbricated plates; secondary radioles numerous, smooth; apical system 
dicyclic. 

Florida species: Species indeterminate and generic identification only 
tentative. 

Comments: Fossils are present in the Chipola Formation of northwestern 
Florida, and represent a new stratigraphic occurrence for the state and a 
potential new taxonomic record. Specimens consist only of plate fragments, thus 
will require much more detailed examination to determinate specific taxonomy. 

cf., Psammechinus sp. 

(Figure 3-14, K) 

Material examined: UF 67463 (figured test fragment). 


117 

Formation: Chipola Formation. 

Locality; Chipola 09 (CA018); Calhoun County, FL; Clarksville 
Quadrangle, SW1/4, Sec. 29, TIN, R9W. 

Collectors: A. Murray and G. Murray. 

Date: 9/30/94. 

Description: One very small test fragment, approximately four mm long 
and wide. Pore pairs circular, closely spaced; poriferous zone in small-scale 
zigzag pattern, with zygopores tracking perimeter of marginal tubercles. Primary 
tubercles with relatively large mamelon, modest scrobicule and scrobicular ring; 
two to three secondary tubercles along perradial suture. 

Comments: Although the preservation of the test fragment is good, the 
overall size greatly limits the identification of the echinoid from which it was 
detached. The general tubercle arrangement and pore pair distribution is similar 
to that of Psammechinus species described from the Eocene of South Carolina 
and the Pliocene of Virginia (Cooke, 1959). Although no Miocene species from 
the U.S. have been reported, this may be a new species that is yet undescribed. 
Until more complete specimens are collected, I can only consider this to be cf. 
Psammechinus sp. for this study. The fossil is included as one of the echinoid 
taxa from the Miocene and a new stratigraphic record for the Chipola Formation 
in the taxonomic total and stratigraphic records tally. 

Unidentified regular echinoid. 

(Figure 3-14, L-M) 

Material examined: UF 1167 (figured internal mold). 


118 

Formation: Arcadia Formation (Tampa Limestone Member). 

Locality: UF locality 2122; Flernando County, FL; near Brooksville, FL 

Collector: Unknown. 

Date: Unknown. 

Description: Florizontal outline circular; aboral surface gently sloping 
dome, slightly flattened near apex. Ambulacra and poriferous zones relatively 
narrow throughout; widest at ambitus, and nearly closing at apical system and 
peristome. Interambulacral plates approximately twice as wide as high; 
interambulacral zone about double the width of ambulacral zone. Pore pairs 
circular; unable to view specific arrangement. Peristome approximately half as 
wide as diameter. Test diameter approximately 39 mm and height 23 mm. 

Comments: Most of the internal mold is poorly preserved, with only 
isolated areas showing any detailed morphology. The general outline, shape, 
and dimensions are similar to species of Arbacia , but the absence of large 
primary tubercle molds on the fossil is not expected, even on an imperfectly 
preserved mold. Due to the limited morphological features visible, I refer to this 
fossil as an unidentified regular echinoid only and will withhold lower taxonomic 
identification until better specimens can be collected. The specimen is counted 
as a new stratigraphic record for the Arcadia Formation, and also is counted as a 
unique species for the Miocene taxonomic list of echinoids. 

Order Clypeasteroida A. Agassiz, 1872 
Family Clypeasteridae L. Agassiz, 1835 
Genus CIvpeaster Lamarck. 1801 


Description: See Oligocene echinoid section for generic description. 


119 


Florida species: C. concavus Cotteau, 1875 and C sp. 

Comments: The species C. concavus is present in the Chipola 
Formation, and the CIvpeaster sp. fossils are present in the Chattahoochee 
Formation. This unidentified species also represents a new stratigraphic record 
of CIvpeaster in the Chattahoochee Formation, and may be a new taxonomic 
record if it is an undescribed species. 

CIvpeaster concavus Cotteau, 1875 
(Figure 3-15, A-B) 

Material examined: UF 65864 (figured test), UF 40318 (test fragment), 
UF 65865 (test), UF 65867 (test). 

Description: Horizontal outline oval to subpentagonal; upper surface 
moderately inflated, petals slightly swollen; lower surface gently rounded near the 
margin, more or less deeply concave around the peristome, ambulacral grooves 
conspicuous; margin rounded. Apical system pentagonal, central; five genital 
pores at the corners of the central madreporite. Petals broad, lanceolate, 
extending about two-thirds the way to the margin, strongly convex near the tips; 
poriferous zones rather broad, completely closed at the apex, nearly closed 
distally; inner pores small, circular; outer pores larger, oval; pores conjugate; 
interporiferous zones more than twice as wide as the poriferous zones at the 
widest part. Peristome central, small. Periproct circular or transversely oval; 
near the margin. Tubercles crowded, sunken, smaller on the upper surface than 
on the lower; one row between each two pairs of zygopores. 


120 

Clypeaster sp. 

(Figure 3-15, C-D) 

Material examined: Eleven internal molds (\A/eathered and incomplete); 
UF 40444 (figured internal mold), UF 66578 (internal mold), UF 66580, UF 
66582, UF 97927. 

Formation: Chattahoochee Formation. 

Locality: Dry Creek 01 (JA010); Jackson County, FL; Oakdale 
Quadrangle, NE1/4, SW1/4, Sec. 11, T3N, R10W. 

Collectors: T. Cassady, B. Shumaker, and C. Shumaker. 

Date: 5/90 (for UF 40444 only). 

Description: Test medium in size; horizontal outline subpentagonal; 
aboral surface relatively flat near ambitus, becoming inflated at apical system. 
Margins relatively thinner than average species of genus, yet thick enough for 
gradual rounding. Adoral surface clearly concave. Apical system located slightly 
anterior of center. Ambulacral petals broadly lanceolate, closed apically and 
slightly open distally; pores circular to subelliptical, conjugate, with each pair 
diverging to maximum at approximate midpoint of petal; outer pore more distal 
than inner pore. Petal length subequal and estimated as two-thirds to three- 
quarters the distance to test margin; no discernible curvature. Tubercles small to 
medium size, relatively densely spaced. Test length of longest specimen 
approximately 55 mm for defined mold only; the overall external test length was 
likely 10-20 mm greater than the internal mold length. 

Comments. All specimens of this species are incomplete internal molds 
with poor to good general preservation. The specimens show some similarity to 


121 


the Miocene species C. concavus (see specific description above), but are 
thinner than C. concavus at the test margin and the petals of the molds are more 
uniform in dimension than those of C. concavus . Therefore, I interpret these 
molds as a new species awaiting formal description, and have included this 
undescribed species as a new stratigraphic record for the Chattahoochee 
Formation as well as a new taxonomic record. 

Gen. et sp. indet. 

(Figure 3-15, E-F) 

Material examined; UF 23086 (figured incomplete test). 

Formation: Coosawhatchie Formation. 

Locality: Brooks Sink (BF001); Bradford County, FL; Brooker 
Quadrangle, SW1/4, SW1/4, Sec. 12, T7S, R20E; east wall of sinkhole, bed 13. 

Collector: R. Portell. 

Date: 1/14/88. 

Description: Single juvenile test, very small (approximately 5 mm 
diameter): central test area missing both apically and adorally. Horizontal outline 
subcircular; widest posterior of center; evidence of initial stages of ambulacral 
notches in ambulacrum I and V; posterior margin somewhat truncated. Test 
margin proportionally medium in thickness, rounded. Both adoral and aboral 
surface flat (although both surface incomplete). Detailed micromorphology not 
visible (i.e., petals, apical system, tubercles, and peristome); periproct positioned 
just posterior of midpoint between peristome and test margin, longitudinally 
elliptical in orientation. 


122 


Comments: The general shape of this fossil is similar to what would be 
expected in a clypeasteroid species, and is similar to other Miocene species such 
as Abertella aberti previously reported from the Coosawhatchie Formation (Jones 
and Portell, 1 988). Since juveniles are not fully developed morphologically and 
this specimen is incomplete, I have chosen to only list its identification to the 
ordinal level at this time. Therefore, I do not include this fossil as a new 
stratigraphic record, new taxonomic record, or as a unique species for the overall 
Miocene diversity value (if it is Abertella aberti . it has already been included from 
this unit in the Miocene). From a biostratigraphic perspective, the only 
significance of this fossil is its presence at the Brooks Sink locality. As work 
continues, additional specimens may be recovered that will provide better 
comparative material for identification purposes, and it may or may not prove to 
be a new species for the Miocene echinoids of Florida. 

Family Fibulariidae Gray, 1855 
Genus Echinocyamus van Phelsum, 1774 

Description; Test moderately flattened; hydropores few, not in groove; 
periproct between first and second pair of coronal plates; petals poorly defined in 
some forms, pore pairs usually oblique; no spicules in tube feet; five pairs of 
internal radiating partitions; in some species females with aboral marsupium. 

Florida species: E. chipolanus Cooke, 1942. 

Comments: This species is present in the Chipola Formation. This is 
one of the rare species reported from Florida that forced me to be solely 
dependent upon previously published information (i.e., Cooke, 1942) as a record 


123 


of its presence in Florida. No samples were personally collected, and no fossils 
are part of the FLMNH collection or other museum and personal collections I 
examined as part of my research. 

Echinocyamus chipolanus Cooke, 1 942 
Material examined: No specimens available in the FLMNFI collection. 
Description: Test small, outline broadly oval, gently arched above, flatter 
below; surface covered with proportionately large tubercles; strengthened by 
internal marginal partitions. Genital pores four in number, widely spaced. 
Ambulacral areas obscure; poriferous zones of each area apparently widely 
diverging. Peristome about one-third the total width, nearly circular. Periproct 
very slightly closer to the margin than to the peristome, rather large, slightly 
pointed posteriorly. 


Family Echinarachniidae Lambert, 1914 
Genus Echinarachnius Gray, 1825 

Description: Petals lyrate, about 0.6 length of radius; periproct marginal, 
between third pair coronal plates; food grooves with straight trunk, two equal 
lateral branches near margin; contact of coronal interambulacral plates with 
primordial plates very variable; posterior area usually discontinuous; three or four 
interambulacral and five or six ambulacral coronal plates on oral surface. 

Florida species: cf. Echinarachnius sp. 

Comments: This tentatively identified genus is present in the Chipola 


Formation. 


124 

cf. Echinarachnius sp. 

(Figure 3-15, G) 

Material examined; UF 67464 (figured test), UF 67465 (test), UF 67483 

(test). 

Formation: Chipola Formation. 

Locality: Chipola 09 (CA018); Calhoun County, FL; Clarksville 
Quadrangle, SW1/4, Sec. 29, TIN, R9W. 

Collectors: A. Murray and G. Murray. 

Date; 9/30/94. 

Description: Test subcircular to subpentagonal; slight invagination along 
margin of ambulacrum I and V; widest point posterior of apical system in 
approximate locations of interambulacrum 1 and 4. Apical system slightly 
inflated; located just anterior of test center. Petaloid ambulacra imperfectly 
developed (all are juveniles); petal length estimated to extend two-thirds the 
distance to margin in ambulacra II, III, and IV, while only half the distance to 
margin in posterior petals. Tubercles proportionally medium size, moderately 
spaced on aboral and adoral surfaces. Peristome slightly eccentric anteriorly; 
medium size (though may have been expanded during compaction and 
preservation), circular in outline. Periproct or possible posterior notch on 
posterior margin; small diameter; elliptical shape. 

Comments: All three specimens are juveniles and clearly not fully 
developed morphologically. This leads to uncertainty in identification of the 
species since allometric growth may produce somewhat different morphological 
proportions in adults than are observed in these fossils. Three genera are 


125 


possible matches for the test characteristics, including Abertella , Protoscutella , 
and Echinarachnius . Of these possibilities, Echinarachnius and Protoscutella are 
the closer matches morphologically, but Protoscutella only is known from the 
Eocene while Echinarachnius has been reported from the Miocene in the western 
U.S. and Japan. Abertella is present in the Miocene of Florida, but these fossils 
are distinctly different in general test shape from Abertella . If these differences 
are simply a reflection of comparison between a juvenile morphology and an 
adult morphology, only collecting additional fossils (that are adults) will allow the 
appropriate comparisons and a solution to this taxonomic question. Therefore, I 
tentatively assign this set of fossils to the genus Echinarachnius and include 
them as a new taxonomic record for the Miocene of Florida and a new 
stratigraphic record for the Chipola Formation. 

Family Mellitidae Stefanini, 1911 
cf. Mellitdae, gen. et sp. incertae. 

(Figure 3-15, H-l) 

Material examined; UF 17635 (figured 2 test fragments). 

Formation: Statenville Formation. 

Locality: Suwannee River Mine (HA002); Hamilton County, FL; 
Benton/Genoa Quadrangles, TIN/1 S, R15/16E. 

Collector: R. Portell. 


Date: 4/9/88. 


126 

Description: Two small, marginal test fragments; surface weathered, 
thereby removing tubercles and micromorphology features. Adoral and aboral 
surfaces flat, terminating in thin, sharp margin, one to two mm thick. 

Comments: Specimens clearly identifiable as derived from flat sand 
dollars, but no identification beyond familial level can be determined. Other 
mellitids are known from the Miocene (see descriptions within this section), but 
until specimens that are more complete are available, the identification will 
remain uncertain. The fossils are included as new stratigraphic records from the 
Statenville Formation, but are not included in the Miocene species diversity total 
due to the limited taxonomic resolution (indeterminable as unique species). 

Family Abertellidae Durham, 1955 
Genus Abertella Durham, 1953 

Description: Petals about 0.7 length of radius; posterior marginal 
indentations most prominent; periproct between second serial pair post- 
basicoronal plates; interambulacra about 0.5 width of ambulacra at ambitus. 

Florida species: Abertella . aberti (Conrad, 1842) and Abertella spp. 

Comments: Fossils of A. aberti are present in the Arcadia and Peace 
River formations. Also reported by Cooke (1959, p. 45) from the Chipola 
Formation along the Sopchoppy River in Wakulla County, but this very likely is an 
inaccurate report, since the Chipola Formation is not present in that area. 
Therefore, I do not include this species as part of the echinoid fauna of the 
Chipola. A second category of fossils awaiting identification are possible 
Abertella species (UF 74785) collected from the Chipola, Torreya, and 


127 


Coosawhatchie formations. These fossils are difficult to identify to genus as well 
as to species, and will require further detailed preparation and examination. The 
specimens have been tentatively identified as possible Protoscutella sp. (B. 
Carter, pers. comm., 1995), but I am not convinced that is the best choice, and 
prefer to include them as an unidentified or undescribed species of Abertella . 

Abertella aberti (Conrad. 1842) 

(Figure 3-16, A-B) 

Material examined: UF 5363 (figured test), UF 104444 (figured test), UF 
14501 (test), UF 25417 (36 test fragments), UF 60072 (test), UF 60073 (test). 

Description: Test shield shaped; horizontal outline semicircular in front, 
fluted behind, with a deep posterior notch of variable width and shallower 
rounded indentations in the posterior ambulacra; upper surface slightly inflated in 
the apical region, nearly flat marginally; oral surface flat; margin thin. Apical 
system fused, star shaped, with the points protruding between the petals and 
four genital pores at the ends, usually somewhat raised. Petals lanceolate, 
extending about two-thirds of the radius; pores small, circular, deeply conjugate, 
inner row nearly straight, outer row broadly arched; poriferous zones wider than 
the interporiferous zones, moderately open at each end. Ambulacra widely 
expanding beyond the petals. Interambulacra narrowing from the outer ends of 
the petals to the margin, slightly expanding within the margin on the oral face but 
interrupted by two long postbasicoronal ambulacral plates. Each submarginal 
plate of the oral side contains a maze of internal passageways. Peristome 
central, small, circular; surrounded by five radiating food grooves, which bifurcate 


128 


at the ends of the basicoronal plates and curve broadly almost to the margin, 
where they branch repeatedly. Periproct on the oral side about three-quarters of 
the radius from the margin, the distance from the margin varying with the depth 
of the posterior notch. 


cf. Abertella sp. 

(Figure 3-16, C-D) 

Material examined: UF 25296 (figured test fragment). 

Formation: Torreya Formation. 

Locality: Gunn Farm Mine (Milwhite Company) (GD006); Gadsden 
County, FL; Dogtown Quadrangle, NE1/4, N1/2, Sec. 74, T4N, R3W. 

Collector: D. Bryant. 

Date: 8/19/88. 

Description: Single test fragment (approximately 30 mm by 39 mm); 
aboral surface flat, sloping gently away from apical system; adoral surface flat. 
Tubercles very small, densely covering both aboral and adoral surfaces. Small 
area of poriferous zone visible; pores very small, circular, conjugate; outer pore 
of pair eccentric marginally relative to inner pore; distal terminus of petal located 
significantly adapically from margin; petal apparently at least partially open at 
end. Test margin interpreted to be thin, possibly sharp. 

Comments: This fossil collected from the Torreya Formation is small, but 
several interpretations can be made regarding its possible taxonomic affinity. 

The specimen clearly is a flat sand dollar variety of clypeasteroid with a thin 
margin. The only Miocene genus known from this age of material with a 


129 


relatively thin test margin is Abertella . A second characteristic of importance is 
the distance the terminal end of the petal lies from the test margin. In species of 
Abertella (such as the Miocene A. aberti), this petal-to-margin distance is at least 
moderate. For example, in many cases it ranges from 10 mm to over 20 mm, 
and in the specimen described above this distance is 17 mm. The original 
complete individual must have been relatively large overall, and species such as 
A. aberti are among the larger clypeasteroid species with respect to test length 
and body size. Therefore, I interpret this fossil as a species awaiting 
identification, but likely a within the genus Abertella . and thus refer to it as cf. 
Abertella sp. It may indeed reflect a new species rather than a new occurrence 
of a previously described species. This genus has been reported from the 
Chipola Formation, but the description herein is a new stratigraphic record from 
the Torreya Formation. I also have counted this as a unique species with regard 
to the overall Miocene diversity, but it is not considered a new taxonomic record 
at this time. 


unidentified Clypeasteroida; cf. Abertella sp. 

(Figure 3-16, E) 

Material examined: UF 23085 (figured test fragment; 70 test fragments 
total in lot). 

Formation: Coosawhatchie Formation. 

Locality: Brooks Sink (BF001); Bradford County, FL; Brooker 
Quadrangle, SW1/4, SW1/4, Sec. 12, T7S, R20E; east wall of sinkhole, bed 13. 


Collector: R. Portell. 


130 


Date: 1/14/88. 

Description; Many very small test fragments (largest dimensions of 14 
mm by 12 mm). Adoral and aboral surfaces flat; tubercles small, circular, closely 
packed on surfaces. Possible marginal fragments thin, sharp. 

Comments: These fossils are similar to fragments of other clypeasteroid 
echinoids found in the Coosawhatchie Formation, particularly Abertella aberti 
fragments. Due to the small size of the fragments, I refer to these fossils as cf. 
Abertella sp., recognizing they may indeed be part of A. aberti individuals known 
to occur in the Coosawhatchie Formation. These fossils are not included in any 
of my diversity or stratigraphic records counts since the generic or specific 
identification is uncertain, and because the most likely species already has been 
verified and included in such tabulations. 

Order Cassiduloida Claus, 1880 
Family Cassidulidae L. Agassiz and Desor, 1847 
Genus Rhvncholampas A. Agassiz, 1869 

Description; See Oligocene echinoid section for generic description. 

Florida species: R. chipolanus Oyen and Portell, 1996, Rhvncholampas 
sp. cf. R. chipolanus . and Rhvncholampas sp. 

Comments: The species R. chipolanus is present in the Chipola 
Formation and the unidentified species of Rhvncholampas is present in the 
Arcadia and Peace River formations. The Rhvncholampas sp. fossils are internal 
molds with relatively poor preservation, thereby making specific identification 
rather difficult. It is possible that these specimens represent new records of R. 


131 


chipolanus from the central part of Florida, but until specimens are collected with 
better preservation, this comparison cannot be completed in detail. Therefore, 
the Rhvncholampas sp. fossils are new stratigraphic records of the genus in both 
the Arcadia and Peace River formations, and possibly a new taxonomic record 
as well. 


Rhvncholampas chipolanus Oyen and Portell, 1996 
(Figure 3-16, F-G) 

Material examined; UF 66633 (figured test). 

Description: Test large, width relatively uniform; margin relatively sharp 
and angular in posterior, subrounded to moderately truncated along lateral 
margins: highest point at apical system, slightly anterior of test center; sides 
slope moderately, with shallower slope in upper half and slightly steeper slope in 
lower half of test. Apical system located central to slightly anterior of test; 
compact size. Petals well developed, broad, lanceolate, with greatest width 
approximately 35 percent of petal length; petals I and V longest. III slightly 
shorter, and II and IV shortest; petals extend approximately 65 percent of 
distance to test margin; petal widths nearly equal, with petal III 25 percent 
narrower than others; petals almost closed at distal end. Periproct 
supramarginal; width approximately 1.5x greater than height; pentagonal shape, 
with apex pointing toward apical system; slight groove extending from periproct 
opening to posterior margin. Peristome moderately anterior, pentagonal, and 
depressed; width approximately 1 .7x greater than length. Floscelle with 
bourrelets distinct and pointed. Adapical tubercles small and uniformly 


132 


distributed: limited visible adoral tubercles near peristome distinctly larger. 
Species characterized by slightly truncated oval outline, moderately sloping sides 
with focused peak at apical system, depressed adoral surface, nearly closed 
lanceolate petaloid ambulacra, and pentagonal periproct. 

Rhyncholampas sp. cf. R. chipolanus Oyen and Portell, 1996. 

(Figure 3-1 7, A-B) 

Material examined: UF 5373 (figured internal mold), UF 10111 (figured 
internal mold). 

Formation: Arcadia Formation. 

Locality: Ft. Green 13 Dragline (PO002); Polk County, FL; Baird 
Quadrangle, Sec. 2, T32S, R23E. 

Collectors D. Jones (UF 5373) and S. King (UF 10111). 

Date: 2/14/87 and 1/86, respectively. 

Description: Test truncated oval in horizontal outline; widest point 
posterior of center; test width/length ratio approximately 0.91 and height/length 
ratio 0.54; aboral surface inflated, somewhat conical and steeply sloping with 
peak anterior of center; adoral surface dominantly obscured but apparently 
slightly depressed. Apical system anterior of center; gonopores not preserved. 
Petaloid ambulacra lanceolate, narrowing at distal end without closing; conjugate 
pore pairs; petals imperfectly preserved but apparently subequal in length, 
extending two-thirds to three-quarters the distance to margin. Periproct 
supramarginal, wider than high, with shallow groove from opening to posterior 
margin; peristome not preserved. 


133 


Comments: I interpret these fossil internal molds to represent a species 
of Rhvncholampas , though imperfect preservation reduces the reliability of 
specific identification. Three described species have similar morphological 
characteristics, including one Miocene species (R. chipolanus ) and two Pliocene 
taxa (R. avresi and R. everqiadensis ). The closest morphological and temporal 
match of the three is R. chipolanus from the Chipola Formation in north Florida 
(see the species description in this section). One aspect of these 
Rhvncholampas fossils that is somewhat unusual is the moldic preservation, and 
this inhibits detailed comparisons with other fossils from the state. Therefore, i 
refer to these specimens as Rhvncholampas sp. cf. R. chipolanus and include 
them as a new stratigraphic record for the Arcadia Formation and as part of the 
diversity total representing R. chipolanus . not a new taxonomic record. 

Rhvncholampas sp. 

(Figure 3-17, C) 

Material examined; UF 12992 (figured test fragment), UF 10671 (test 
fragment). 

Formation: Peace River Formation. 

Locality: Zolfo Springs (FIR001); Flardee County, FL; Zolfo Springs 
Quadrangle, Sec. 27/28, T34S, R25E. 

Collector; C. Hewlett. 

Date; Unknown. 

Description: Test fragmented, weathered, and incomplete; interpreted as 
relatively large. Aboral surface inflated, broadly curving: moderately tall. Apical 


134 


system destroyed. Isolated remnants of petaloid ambulacra present; conjugate 
pore pairs, circular; outer and inner pores diverging at maximum width of petal, 
outer pore eccentric marginally as compared with inner pore of pair; pores 
tightening distally and presumed similar proximally; petals lanceolate, not 
completely closed at distal end. Adoral surface dominantly flat with slight 
concavity near peristome. Peristome damaged but located anterior of center; 
bourrelets present; periproct destroyed. Tubercles small, closely spaced both 
adorally and aborally. 

Comments: This fossil is significantly damaged, thereby preventing full 
identification. However, the overall shape of the remaining portion is identifiable 
as a species of Rhvncholampas . The Miocene of Florida only has one species 
reported (R. chipolanus Oyen and Portell, 1996 from the Chipola Formation; see 
description in this section), and this specimen may be a new record of C. 
chipolanus for the Peace River Formation. Without a better preserved specimen, 
it is impossible to fully compare with other species and must be left as species 
indeterminate. Herein I consider this fossil as a new stratigraphic record for the 
Peace River Formation (as the first stratigraphic record of Rhvncholampas in the 
unit), but do not include it as a new taxonomic record for the Miocene nor as part 
of the diversity total for the epoch. 

Order Spatangoida Claus, 1876 
Family Schizasteridae Lambert, 1905 
Genus Aqassizia L. Agassiz and Desor, 1847 


Description: See Eocene echinoid section for generic description. 


135 

Florida species; Aqassizia sp. 

Comments: The species is present in the Arcadia Formation (Miocene), 
from the Dean’s Trucking Pit locality in southern Florida. This represents a new 
stratigraphic record, and likely a new taxonomic record for Aqassizia in the 
Miocene (since no Miocene species are known from the southeastern U.S.). 

Aqassizia sp. 

(Figure 3-17, D) 

Material examined; UF 28401 (figured incomplete internal mold). 

Formation: Arcadia Formation. 

Locality; Dean’s Trucking Pit (SO006); Sarasota County, FL; Laurel 
Quadrangle, SW1/4, Sec. 22, T38S, R19E. 

Collectors: R. Portell and P. Whisler. 

Date: 8/16/86. 

Description; Internal mold, incomplete and compacted. Horizontal 
outline interpreted as subcircular to subovate. Aboral surface inflated, 
hemispherical-shaped, sloping steeply toward margins; margins broadly rounded; 
adoral surface not present in sample. Apical system posterior of center; plate 
arrangement not preserved. Ambulacra slightly depressed; ambulacrum III 
poorly developed; ambulacra II and IV longest, extending two-thirds the distance 
to margin; anterior paired ambulacra diverging at approximately 40-45° from 
anterior-posterior medial line at proximal end, curving slightly toward posterior at 
distal end; posterior ambulacra pair shorter, slightly wider, and approximately half 
the length of anterior pair, and diverging at about 40-45° from medial line. Entire 


136 


adoral surface absent; posterior margin dominantly crushed or absent. Overall 
test size small with test length approximately 13 mm and width approximately 14 
mm. 

Comments; Although this fossil is poorly preserved, the size, shape, and 
petaloid ambulacra support the identification as a species of Aqassizia . The 
genus has been reported from the Eocene (A. clevei) . Oligocene (A. mossomi) 
and Pliocene (A. porifera ) of Florida, and with this description is a new record 
from the Miocene. The preservation is too poor to define the specific level 
identification, and therefore I refer to the fossil only as Aqassizia sp. It repre- 
sents a new stratigraphic record from the Arcadia Formation, and likely is a new 
taxonomic record (and counted as such herein) since no other Miocene species 
are known from the Gulf and Atlantic Coastal Plains of the southeastern U.S. 

Family Brissidae Gray, 1855 
Genus Brissopataqus Cotteau, 1863 

Description; Differs from Eupataqus in having large depressions in front 
of anterior petals or in front of all petals. 

Florida species; Brissopataqus sp. 

Comments; The species is present in the Chattahoochee Formation 
along Dry Creek in northern Florida. 

Brissopataqus sp. 

(Figure 3-18, A-B) 

Material examined; UF 97921 (figured internal mold), UF 40441 (figured 
internal mold), UF 66568 through 66577 (one internal mold each). 


137 


Formation: Chattahoochee Formation. 

Locality: Dry Creek 01 (JA010); Jackson County, FL; Oakdale 
Quadrangle, NE1/4, SW1/4, Sec. 11, T3N, R10W. 

Collectors: T. Cassady, B. Shumaker, and C. Shumaker. 

Date: 5/90. 

Description: Horizontal outline asymmetrically oval; narrov\/est and 
truncated posteriorly, widest slightly anterior of center. Aboral surface broadly 
curving at margin, somewhat elevated, and flattening near center of test; apical 
system anteriorly eccentric, at one-third the length from anterior margin. Petaloid 
ambulacra shallowly depressed; anterior pair (II and IV) diverge at about 160°; 
posterior pair (I and V) diverge at about 50°; ambulacrum III more poorly 
developed, flush with surface, leading into small anterior sulcus at test margin. 
Petals curve slightly toward anterior at distal ends; anterior pair curvature more 
pronounced than posterior pair; posterior petals shorter than anterior pair, 
extending approximately half the distance to margin; anterior pair extend three- 
quarters the distance (or slightly more) to margin. Petals lanceolate, narrowing 
minimally at distal end yet remaining open. Zygopores circular, moderately 
spaced; outer pore slightly advanced toward margin. Adoral surface moderately 
flattened; locally depressed around peristome, and raised along a medial ridge 
extending from peristome to posterior margin. Peristome at anterior quarter; 
width approximately double the length; curved toward posterior; labrum not 
conspicuously overhanging. Periproct inframarginal to supramarginal at posterior 
truncation (rarely preserved well enough to describe). Tubercles not preserved. 


138 


Comments. These internal molds are moderately well preserved, 
allowing identification to the generic level of Brissopataqus . One species in this 
genus was reported by Checchia-Rispoli (1947) from Oligocene-Miocene rocks in 
northern Africa. However, no species have been reported from the Miocene of 
the Caribbean or the southeastern U.S. and, therefore, it is likely these fossils are 
a new species. Thus, I refer to these fossils as Brissopataqus so. herein and 
include them as a new stratigraphic record for the Chattahoochee Formation, a 
new taxonomic record for the genus, and count this unnamed species in my 
diversity total for the Miocene of Florida. 

cf. Brissidae; gen. et sp. indet. 

(Figure 3-18, C-D) 

Material examined: UF 67379 (figured test fragment), UF 44622 (figured 
test fragment). 

Formation; Chipola Formation. 

Locality; UF 67379: Farley Creek 07 (CA002); Calhoun County, FL; 
Clarksville Quadrangle, SW1/4, NE1/4, Sec. 21, TIN, R9W. UF 44622: Tenmile 
Creek 02 (CA003); Calhoun County, FL; Altha West Quadrangle, N1/2, Sec. 

11/12, TIN, R10W. 

Collectors; R. Portell (UF 67379) and Brooks, Scolaro, and Dimmick (UF 
44622). 

Date; 4/29/94 and 11/9/73, respectively. 

Description: The single test fragment (UF 67379) is a coherent plastron 
from posterior adoral surface. Ridge reduced, peak near posterior margin of 


139 


plate; small tubercles of fascicle distributed throughout, largest near plate 
margins. Second lot (UF 44622) dominated by very small plate fragments. 
Fragments covered by closely spaced tubercles; tubercles with pronounced, 
cylindrical boss, distinct parapet, and enlarged, perforate mamelon. 

Comments; This set of fossils is so limited with respect to overall test 
morphology that only general statements regarding identification can be 
provided. The single plastron is similar to features found in various species of 
the family Brissidae in the order Spatangoida. Relatively few species of brissids 
have been reported from Miocene units in the U.S. and the Caribbean. One 
brissid was described and named from a possible Miocene formation in South 
Carolina by Cooke in 1959, that he named Soatangus glenni Cooke, but has 
since been revised to Brissus glenni (Cooke). The plastron of Cooke’s species is 
morphologically similar to the one reported herein from the Chipola Formation, 
and may be closely related or possibly the same taxon. The second group of test 
fragments (UF 44622) are not distinct taxonomically, but are the appropriate size 
to have been part of one of the fascioles on a brissid (perhaps the subanal 
fasciole or the marginal fasciole). Specimens that are significantly more 
complete are required before any lower taxonomic identifications can be 
proposed for these fossils. The fossils are included as a new stratigraphic record 
for the Chipola Formation (although previously mentioned in a non-specific 
manner in Oyen and Portell, 1996, p. 60), as a possible new taxonomic record, 
and as a unique species for the Miocene diversity total. 


Figure 3-14. Miocene regular echinoids. 

A) Prionocidaris cookei Cutress, 1976; UF 66632; lateral view of test fragment; 
Chipola Formation; 1.5x. 

B) Prionocidaris cookei Cutress. 1976; UF 88540; four incomplete radioles; 
Chipola Formation; 1.5x. 

C) Prionocidaris cookei Cutress. 1976; UF 101422; exterior of three 
disarticulated, imperforate test plates; Chipola Formation; 1.5x. 

D) CIDAROIDA, gen. et sp. indet.; UF 66579; lateral view of dolomitized, partial 
internal mold of test; Chattahoochee Formation; lx. 

E) CIDAROIDA; gen. et. sp. indet.; UF 66513; lateral view of incomplete radiole; 
Shoal River Formation; 2.x. 

F) Arbia sp.; UF 102309; lateral view of dolomitized, partial external mold of test; 
Chattahoochee Formation; lx. 

G) Arbia sp.; UF 102309; lateral view of RTV silicone rubber peel made from 
partial external mold mentioned above; lx. 

FI) Gaqaria sp.; UF 25339; lateral view of test fragment; Parachucia Formation; 
2x. 

I) cf. Gaqaria sp.; UF 60668; aboral view of internal mold of test; Chattahoochee 

Formation; lx. 

J) cf. Gaqaria sp.; UF 60668; adoral view of internal mold of test; Chattahoochee 
Formation; lx. 

K) cf. Psammechinus sp.; UF 67463; lateral view of test fragment; Chipola 
Formation; 3x. 

L) Unidentified regular echinoid; UF 1167; aboral view of internal mold of test; 
Arcadia Formation; lx. 

M) Unidentified regular echinoid; UF 1167; adoral view of internal mold of test; 
Arcadia Formation; lx. 


141 





Figure 3-15. Miocene irregular echinoids. 

A) CIvpeaster concavus Cotteau, 1875; UF 65864; aboral view of test; Chipola 
Formation; lx. 

B) CIvpeaster concavus Cotteau, 1875; UF 65864; adoral view of test; Chipola 
Formation; lx. 

C) CIvpeaster sp.; UF 40444; aboral view of dolomitized, partial test; 
Chattahoochee Formation; lx. 

D) CIvpeaster sp.; UF 66578; adoral view of dolomitized, partial test; 
Chattahoochee Formation; lx. 

E) Fam., gen., et sp. indet.; UF 23086; aboral view of incomplete, juvenile test; 
Chipola Formation; 3x. 

F) Fam., gen., et sp. indet.; UF 23086; adoral view of incomplete, juvenile test; 
Chipola Formation; 3x. 

G) cf. Echinarachnius sp.; UF 67464; aboral view of juvenile test; Chipola 
Formation; 3x. 

FI) cf. Mellitidae; UF 17635; test fragment; Statenville Formation; 2x. 

I) cf. Mellitidae; UF 17635; test fragment; Statenville Formation; 2x. 


143 





Figure 3-16. Miocene irregular echinoids. 

A) Abertella aberti (Conrad. 1842); UF 5363; aboral view of test; Arcadia 
Formation; 0.75x. 

B) Abertella aberti (Conrad. 1842); UF 104444; adoral view of test; Arcadia 
Formation; 0.75x. 

C) cf. Abertella sp.; UF 25296; aboral view of test fragment; Torreya Formation; 
lx. 

D) cf. Abertella sp.; UF 25296; adoral view of test fragment; Torreya Formation; 
lx. 

E) Unidentified Clypeasteroidea; cf. Abertella sp.; UF 23085; test fragment; 
Coosawhatchie Formation; 2x. 

F) Rhvncholampas chipolanus Oyen and Portell, 1996; UF 66633; aboral view of 
test; Chipola Formation; lx. 

G) Rhvncholampas chipolanus Oyen and Portell, 1996; UF 66633; adoral view 
of test; Chipola Formation; lx. 

H) Rhvncholampas chipolanus Oyen and Portell, 1996; UF 66633; lateral view of 
test; Chipola Formation; lx. 


145 




Figure 3-17. Miocene irregular echinoids. 

A) Rhvncholampas sp. cf. R. chipolanus Oyen and Portell, 1996; UF 5373; 
aboral view of dolomitized, incomplete internal mold of test; Arcadia 
Formation; lx. 

B) Rhvncholampas sp. cf. R. chipolanus Oyen and Portell, 1 996; UF 10111; 
aboral view of dolomitized, incomplete internal mold of test; Arcadia 
Formation; lx. 

C) Rhvncholampas sp.; UF 12992; lateral view of incomplete test embedded in 
sandstone matrix; Peace River Formation; lx. 

D) Aaassizia sp.; UF 28401; aboral view of dolomitized, incomplete internal mold 
of test; Arcadia Formation; 2x. 


147 




Figure 3-18. Miocene irregular echinoids. 

A) Brissopataqus sp.; UF 97921 ; aboral view of dolomitized, internal mold of 
test; Chattahoochee Formation; lx. 

B) Brissopataqus sp.; UF 40441; adoral view of dolomitized, internal mold of 
test; Chattahoochee Formation; lx. 

C) cf. Brissidae ; gen. et sp. indet.; UF 67379; adoral view of test plastron; 
Chipola Formation; lx. 

D) cf. Brissidae ; gen. et sp. indet.; UF 44662; test fragments; Chipola Formation; 
2x. 

E) Lovenia clarki (Lambert, 1 924); UF 61083; aboral view of internal mold of 
test; Chattahoochee Formation; lx. 

F) Lovenia clarki (Lambert, 1924); UF 61083; adoral view of internal mold of test; 
Chattahoochee Formation; lx. 

G) Lovenia clarki (Lambert, 1 924); UF 61083; adoral view of external mold of 
test; Chattahoochee Formation; lx. 

FI) Lovenia clarki (Lambert, 1 924); UF 61083; RTV silicone rubber peel of adoral 
view of internal mold of test; lx. 


149 



»>»/j 


Family Loveniidae Lambert, 1905 
Genus Lovenia Desor, 1847 


150 


Description: Test low, oval to heart-shaped, with subanal and internal 
fascioles; three or four gonopores; spheridia housed in cysts surrounding 
peristome; in some species primary tubercles of paired ambulacral areas 
recessed into camellae. 

Florida species: L clarki (Lambert, 1924). 

Comments: The species is present in the Chattahoochee Formation. 
Thus far, only the Jim Woodruff Dam locality in northern Florida has been 
reported to have these fossils, but they are relatively common and moderately 
preserved at this site as external and internal molds. 

Lovenia clarki (Lambert, 1924) 

(Figure 3-18, E-H) 

Material examined: UF 61083 (figured internal and external molds with 
RTV silicone rubber peels), UF 66934 through 66939 (one internal mold each). 

Description: Test cordate, depressed, truncated behind. Known only as 
molds, which show cross-shaped petals, evidently terminated apically by an 
apical fascicle. 

Pliocene Echinoids 

Order Cidaroida Claus, 1880 

Genus Eucidaris Pomel, 1883 

Description: Like Stviocidaris but madreporite slightly larger than other 
genital plates; primary spines typically cylindrical, truncate, otherwise fusiform or 


151 


clavate; shaft abruptly truncate, terminating in crown with central prominence, 
and with low, rounded warts disposed in regular, longitudinal series; secondary 
spines adpressed; tridentate pedicellariae of two types, valves either straight or 
curved. 

Florida species; E. tribuloides (Lamarck, 1816). 

Comments: The species occurs in the Tamiami Formation in 
southwestern Florida (see Portell and Oyen, 1997, for detailed discussion). 

Eucidaris tribuloides (Lamarck. 1816) 

(Figure 3-19, A-B) 

Material examined: UF 72022 (figured test), UF 60203 (test), UF 80300 
(radicle), UF 68891 (test with spines), UF 62735 (35 radicles). 

Description: Apical system subcircular or subpentagonal, easily 
detached, less than half the horizontal diameter. Periproct pentagonal, covered 
with polygonal plates. Peristome circular, nearly half the horizontal diameter, 
covered with imbricating plates. Ambulacra nearly straight; poriferous zones 
narrower than the interporiferous zones; zygopores slightly oblique, pores 
separated by a projecting wall; interporiferous zones bordered on each side by a 
row of imperforate miliary tubercles interspersed with small granules. 
Interambulacra composed of five to twelve tiers of plates. Each plate supports 
on its outer edge one large raised perforated tubercle surrounded by a ring of 
imperforate miliary tubercles; median part covered by transverse rows of smaller 
imperforate miliary tubercles. Primary tubercles supporting long, rather thick, 
cylindrical spines decorated with longitudinal rows of circular nodes; miliary 


152 


circles bearing short, flat, ribbed spines; other miliary tubercles bearing shorter, 
narrower, flat spines. 


Order Arbacioida Gregory, 1900 
Family Arbaciidae Gray, 1855 
Genus Arbacia Gray, 1835 

Description; Test low hemispherical or subconical, flattened adorally, of 
medium size. Ambulacra with trigeminate plates, pore zones straight, narrow 
above ambitus, conspicuously widened adorally. Primary ambulacra tubercles in 
regular series. Interambulacra with numerous primary tubercles in horizontal and 
vertical series. No secondary tubercles. Adapically interambulacra have 
conspicuous naked spaces. 

Florida species: A. improcera (Conrad, 1843) and a second possible 
Arbacia sp. 

Comments: Collected from the Tamiami and Jackson Bluff formations. 
Occurrence in the Jackson Bluff Formation represents new stratigraphic record, 
while the Arbacia sp. represents a new taxonomic record when it is formally 
described in the future. Due to the poor preservation of this fossil, it is very 
difficult to determine its specific status at the current time. 

Arbacia improcera (Conrad, 1843) 

(Figure 3-19, C) 

Material examined: UF 83420 (figured test), UF 21399 (test with spines), 


UF 30406 (test), UF 47129 (test). 


153 

Description: Test large, horizontal outline circular; upper surface 
significantly depressed; lower surface evenly rounded and concave. Apical 
system dicyclic; plates covered with elongated granules; ocular plates having one 
high imperforate tubercle. Ambulacra narrow, regularly expanding to the 
ambitus, maintaining nearly the maximum width to the peristome; poriferous 
zones straight on upper surface, expanding near the peristome because of the 
increasing inclination of the groups of three zygopores; one spheridial pit in the 
middle of each area near the peristome. Interambulacral plates nearly three 
times as wide as high, tubercle-free areas covered by coarse, elongate granules. 
Periproct rather large, oblique. Tubercles high, imperforate; largest on margin 
and lower surface; two rows in each ambulacrum, becoming obsolete toward the 
apex; four rows on and below the ambitus in each interambulacrum, only the 
outer rows, much reduced in size, continuing to the apex. Peristome large, 
subpentagonal. 


Arbacia sp. 

(Figure 3-19, D-E) 

Material examined; UF 7506 (figured test, nearly complete, although 
partially obscured by epitaxial cement and matrix). 

Formation: Jackson Bluff Formation. 

Locality; Jackson Bluff 02 (LN002), Leon County, Florida, Bloxham 
Quadrangle, Sec. 16, SW 1/4, T1S, R4W. 

Collector: N. Weisbord. 


Date: Unknown. 


154 


Description: Test small; diameter approximate 17 mm; horizontal outline 
circular and upper surface flattened to only slightly domed; apical system plates 
missing; adoral surface shows evidence of concavity toward peristome, but most 
of surface covered with tightly cemented matrix. Ambulacra relatively straight, 
narrow, expanding evenly to ambitus; zygopores possibly nonconjugate. 
Interambulacral plates approximately two to three times as wide as tall. 

Tubercles small, in two rows in interambulacrum, slightly increasing in size at 
ambitus and adoral surface; one to two additional rows of tubercles present at 
ambitus and below. Periproct absent (due to diagenetic compaction); peristome 
not visible (covered by matrix). 

Comments: This fossil compares reasonably well with the characteristics 
of Arbacia, but the specific identification has not been determined yet. Based on 
the characteristics listed above, it appears to be different from another Florida 
species, A. improcera . However, it may be more closely related to an extant 
species, A. punctulata (Lamarck, 1816), than other described fossil species. 
Unfortunately, only one specimen has been collected thus far and it is not 
preserved well enough to allow identification with certainty. 

Arbacia sp. cf. A. improcera (Conrad, 1843) 

(Figure 3-19, F) 

Material examined: UF 7507 (figured test fragment). 

Formation: Jackson Bluff Formation. 

Locality: Jackson Bluff 02 (LN002), Leon County, Florida, Bloxham 
Quadrangle, Sec. 16, SW 1/4, T1S, R4W. 


155 


Collector: N. Weisbord. 

Date: Unknown. 

Description: Test incomplete, with diameter estimated at 35-40 mm; 
horizontal outline likely circular and upper surface moderately arched. 

Ambulacra straight, reaching maximum width at ambitus; pore pairs closely 
spaced, conjugate, circular in outline. Interambulacral plates approximately three 
times as wide as tall; non-granular near apex. Tubercles largest at ambitus and 
below, decreasing in size adapically; distribution somewhat irregular moving 
toward apical system, more abundant at ambitus and lower surface. Periproct 
and peristome not present on test fragment. 

Comments: The test is significantly fragmented and incomplete 
(estimated to be only 30% of original total), thereby limiting interpretations and 
identification. Preservation of the remaining material is very good. This 
specimen is different from the other regular echinoid (which I refer to as Arbacia 
sp.) reported herein from the Jackson Bluff Formation, and therefore the number 
of taxa from this unit reflects this interpretation. One of the closest matches for 
this fossil is another Pliocene species, A. improcera , that is present in the 
Tamiami Formation in Florida. Since the identification is tentative, this fossil 
does not represent a new taxonomic record for the Pliocene but is a new 
stratigraphic record of the genus for the Jackson Bluff Formation, regardless of 
specific identification. 


156 

cf. Arbacia sp. 

(Figure 3-19, G) 

Material examined: UF 84283 (figured test fragment). 

Formation; Nashua Formation. 

Locality: Cracker Swamp Ranch 01 (PU004); Putnam County, FL; 
Hastings Quadrangle, SE1/4, SW1/4, Sec. 24, T9S, R27E. 

Collectors: R. Portell and C. Oyen. 

Date; 6/26/96. 

Description: Test fragment small, approximately 18 mm by 12 mm. 
Interpreted shell shape subhemispherical, curving moderately at ambitus. 
Tubercles largest at ambitus, becoming distinctly smaller adapically; tubercles 
with large boss and globose mamelon, scrobiculate with well-defined scrobicular 
ring. Poriferous zone consisting of two nearly straight columns of zygopores; 
three to four pore pairs per ambulacral plate. Interambulacral plates becoming 
granular adapically. Ambulacral areas approximately two thirds as wide as 
interambulacral regions. 

Comments: The test fragment was derived from the ambitus of the 
original echinoid. Using the tubercle size and distribution in this area as a 
primary morphology guide, I interpret this pattern as similar to species of Arbacia . 
Therefore, I refer to this specimen as cf. Arbacia sp., and include it as a new stra- 
tigraphic record of the genus from the Nashua Formation but do not include it as 
a new taxonomic record or as a unique species for the Pliocene diversity count. 


157 


Order Temnopleuroida Mortensen, 1942 
Family Toxopheustidae Troschel, 1872 
Genus Lytechinus A. Agassiz, 1863 

Description: Medium-sized to large, low hemispherical. Ambulacra 
plates trigeminate, each with primary tubercle; secondary ambulacra tubercles 
not in regular series; conspicuous naked median space aborally in both areas. 
Buccal membrane bearing numerous plates, in addition to oral plates. 

Florida species: L. varieqatus plurituberculatus Kier, 1963. 

Comments: This fossil is present in the Tamiami and Caloosahatchee 
formations in southern Florida. 

Lytechinus yarieqatus plurituberculatus Kier, 1 963 
(Figure 3-19, H-l) 

Material examined: UF 12895 (figured test), UF 25764 (test), UF 30732 
(test), UF 62805 (test fragment), UF 64825 (test). 

Description: Test hemispherical, large. Apical system with serrate 
outline; the two posterior ocular plates insert. Periproct large, oyal, eccentric. 
Ambulacra wide, trigeminate, the upper two zygopores of each plate near the 
outer edge, the lower one near the tubercle; zygopores forming a continuous row 
along the peristome in each ambulacrum. Plates much wider than high near the 
ambitus. Tubercles smooth, imperforate; two continuous rows in each area, 
passing through the center of each plate; additional rows in the interambulacra 
not reaching the apex, leaying bare median areas. Peristome subpentagonal; 
about one-third the total diameter; gill slits deep. Spines rather short, tapering. 


158 

longitudinally ribbed. Distinguished from nominate subspecies by more 
numerous tubercles in ambulacra. 

Family Echinometridae Gray, 1825 
Genus Echinometra Gray, 1825 

Description; Ambitus oblong or elliptical, longer transverse axis passing 
through ocular I and genital 3; ambulacral plates polyporous, quadrageminate to 
decageminate, exceptionally trigeminate; spines equal to or shorter than test 
diameter, acuminate but not otherwise modified. 

Florida species: E. lucunter (Linnaeus. 1758). 

Comments; The species is present in the Caloosahatchee Formation of 
southern Florida. Fossils consist of test fragments and spines. 

Echinometra lucunter (Linnaeus, 1758) 

(Figure 3-19, J-K) 

Material examined; UF 12937 (figured test), UF 27516 (test), UF 64824 

(test). 

Description; Horizontal outline obliquely elliptical or subpentagonal; 
variably inflated; ambitus broadly rounded. Apical system rather large; one or 
more ocular plates insert or not; madreporite swollen. Periproct oval, slightly 
eccentric. Ambulacra regularly expanding to the ambitus, about equal in width to 
the interambulacra at the peristome; zygopores arranged in disconnected arcs of 
six, more or less, except near the peristome, where they lie in straight, oblique 


159 

lines. Peristome nearly circular, weakly notched. Primary tubercles large, 
smooth, imperforate; in two rows on each area, largest on the interambulacra. 

Order Clypeasteroida A. Agassiz, 1872 
Family Clypeasteridae L. Agassiz, 1835 
Genus CIvpeaster Lamarck, 1801 

Description; See Oligocene echinoid section for generic description. 

Florida species: Up to seven species have been collected in the state, 
including C. crassus Kier, 1 963, C. rosaceus dalli (Twitchell, 1915), C. 
subdepressus (Gray, 1825), C. sunnilandensis Kier, 1963, and three possible 
new species referred to herein as CIvpeaster sp. 

Comments; The species C. crassus . C. sunnilandensis , and CIvpeaster 
sp. are present in the Tamiami Formation. The species C. rosaceus dalli and C. 
subdepressus are present in the Caloosahatchee Formation. The second 
unidentified species of CIvpeaster is present in the Intracoastal Formation, which 
is a new stratigraphic record of the genus from the Intracoastal. The final 
unidentified species of CIvpeaster is present in the Nashua Formation. This 
occurrence is a new stratigraphic record of the genus in the Nashua. Each 
CIvpeaster sp. groups is a potential new taxonomic record for the Pliocene as 
well. 

CIvpeaster crassus Kier, 1963 

Material examined: None available for examination in the FLMNFI 


collection. 


160 


Description: Average test width 90 percent of length, average height 19 
percent of length; test pentagonal with truncated posterior margin, pointed 
anterior with greatest width anterior to center; strong indentations in 
interambulacra 1, 4, 5; margin thick, 10 percent of length, area between margin 
and ends of petals flat or slightly depressed; petaloid area inflated; adoral surface 
flat. Apical system slightly posterior to center, five genital pores, small ocular 
plates, madreporite star-shaped. Petals broad, short, extending three-fifths 
distance from apical system to margin; anterior petal (III) slightly longer than 
others, anterior paired petals (II, IV) shortest, posterior paired petals (V, I) 
intermediate; interporiferous zone approximately twice width of poriferous zone; 
approximately 60 pore pairs in each poriferous zone. Periproct inframarginal, 
located near posterior margin; opening irregular in outline, elongated 
transversely. Peristome central to slightly posterior, pentagonal, pointed 
anteriorly, truncated posteriorly. Plate sutures of basicoronal plates not visible on 
all plates; basicoronal interambulacral plates separated from postbasicoronal 
plates by two pairs of ambulacral plates; seven to eight ambulacral, three to five 
interambulacral postbasicoronal plates in each series on adoral surface. Species 
characterized by thick margin and marginally indented interambulacra. 

CIvpeaster sunnilandensis Kier, 1 963 
(Figure 3-20, A-B) 

Material examined; UF 22148 (figured test). 

Description; Fossils large; test elongate, average width 85 percent of 
length; marginal outline pentagonal, anterior pointed, posterior truncated. 


161 


interambulacra 4 and 1 slightly indented at margin; area between margin and 
ends of petals sloping marginally; test low, average height 20 percent of length; 
margin thin, thickness approximately seven percent of length; petaloid area 
inflated, adoral surface slightly depressed. Apical system central to slightly 
anterior, five genital pores, small ocular plates, madreporite star-shaped. Petals 
broad, of unequal length, anterior petal (III) longest, 20 percent longer than 
anterior paired petals (II, IV); posterior paired petals intermediate in length; 
anterior petal open, gap at distal end of petal 4.4 percent of length, posterior 
petals open in some specimens; interporiferous zone approximately twice width 
of poriferous zone. Periproct inframarginal, located near posterior margin, 
opening irregular in outline, elongated transversely. Peristome central. Plate 
sutures not visible on any specimens. Species characterized by thick margin and 
marginally indented interambulacra. 

CIvpeaster rosaceus dalli (Twitchell. 1915) 

(Figure 3-21, A-B) 

Material examined; UF 65813 (figured test), UF 62796 (2 tests), UF 
62797 (2 tests). 

Description; Test large; pentagonal in marginal outline, longer than 
broad, broadest opposite the ends of the anterior petals, pointed anteriorly, 
posterior end truncated centrally; upper surface irregularly convex, high, highest 
back of center, whence sloping gently and in a straight line to the blunt, tumid 
anterior edge and quite steeply and in a straight line to the thinner, wedge- 
shaped posterior edge; under surface deeply concave, the concavity beginning 


162 


near the margin and increasing at first gradually then rapidly to the center. 
Ambulacral petals large, broad, very tumid, almost reaching the margin, nearly 
closing, the posterior pair slightly longer than the odd petal which is slightly 
longer than the anterior pair. Apical system central, sloping downward anteriorly; 
five genital pores a short distance from the slightly depressed madreporite. 
Peristome large, slightly eccentric posteriorly, subpentagonal, deeply sunken; 
ambulacral furrows simple, straight, reaching the margin. Periproct rather large, 
subcircular; inframarginal, almost marginal. 

CIvpeaster subdepressus (Gray, 1825) 

(Figure 3-22, A-C) 

Material examined: UF 98692 (figured test), UF 21532 (figured Recent 
test), UF 49994 (test), UF 49995 (test), UF 67249 (test). 

Description: Test large; horizontal outline subovate to subpentagonal, 
usually truncate or reentrant in the interambulacra; upper surface tumid in the 
petaloidal region, flatter marginally; lower surface nearly flat, slightly concave at 
the peristome, having narrow ambulacral furrows, the paired furrows bent 
towards each other near the peristome; margin rather thin. Apical system 
pentagonal, with five genital pores at the corners of the madreporite. Petals 
rather short, extending more than halfway to the margin, the anterior the longest; 
poriferous zones curved inward at the outer tips but more or less open, pores 
near the apex inconspicuous, thereby seemingly wide open; interporiferous 
zones wide; pores conjugate. Peristome central, circular. Periproct near the 
posterior margin. Tubercles small. 


CIvpeaster sp. 
(Figure 3-23, A) 


163 


Material examined: UF 104521 (figured incomplete test). 

Formation: Intracoastal Formation. 

Locality: Pickett Bay 01 (FR001); Franklin County, FL; Pickett Bay 
Quadrangle, NW1/4, NW1/4, Sec. 34, T6S, R5W. 

Collectors: R. Portell and K. Schindler. 

Date: 5/6/93. 

Description: Horizontal outline pentagonal, gently rounded at anterior 
point; test margin medium to thin, rounded; test medium to large in dimension. 
Aboral surface covered with tightly cemented matrix and not visible. Adoral 
surface nearly flat, with minor concavity toward peristome; narrow, straight 
ambulacral furrows from margin to peristome. Peristome small, circular, located 
slightly posterior of center. Periproct circular to subcircular, small, near posterior 
margin. Tubercles small, uniformly distributed across test. 

Comments: Only the adoral surface is visible in the described specimen, 
thus limiting full identification. The aboral surface is heavily coated with strongly 
cemented matrix, and this preservation style (i.e., surficially cemented matrix) 
affects most of the samples associated with the Intracoastal Formation from this 
locality. As a result, most of the samples are very difficult to clean, prepare, and 
consequently identify. The general test size, shape, and oral surface 
characteristics are consistent with those of the genus CIvpeaster and therefore I 
refer to this fossil as CIvpeaster sp. A species from the Caloosahatchee 
Formation, C. subdepressus , has a similar morphology for the adoral surface, but 


164 


without the diagnostic characteristic of the aboral surface to compare, I have 
chosen to defer a specific identification until better fossils are recovered. This 
fossil is the first report of the genus from the Intracoastal Formation and therefore 
is counted as a new stratigraphic record from the formation, but is not included 
as an additional species in the total species diversity from the Pliocene epoch. 

CIvpeaster sp. 

(Figure 3-23, B-E) 

Material examined; UF 30870 (figured test fragment), UF 44202 (figured 
test fragment), UF 35666 (test fragment). 

Formation; Tamiami Formation. 

Locality; Richardson Road Shell Pit 01 (SO019), Sarasota County, FL; 
Bee Ridge Quadrangle, Sec. 7/8, T36S, R19E; Phase 1 in pit, spoil samples 
(both UF 35666 and UF 30870). Macasphalt Shell Pit (SO001), Sarasota 
County, FL; Bee Ridge Quadrangle, E 1/2, Sec. 12, T36S, R18E; spoil sample 
(UF 44202). 

Collectors; R. Portell (UF 30870), D. Bryant (UF 44202), and K. 

Schindler (UF 35666). 

Date; 8/19/89, 3/21/87, 12/08/90, respectively. 

Description; Qnly three test fragments total have been collected and 
were used for this description. Test margin broadly rounded. Test thins toward 
margin and gently thickens adapically. Plates thick in cross-section. Tubercles 
small, closely spaced, and densely distributed both on adoral and aboral 
surfaces. No diagnostic morphological features, including peristome, periproct. 


165 


zygopores, gonopores, apical system, petaloid ambulacra, present in these fossil 
fragments. Modest food groove furrow noted on adoral surface of one fossil. 

Test thickness at margin ranges from approximately four to seven mm. 

Comments; Although only limited fragments of this echinoid taxon have 
been collected to date, identification to generic level is reasonably valid. The 
overall shape of the test margin, its thickness, and the tubercle density tends to 
support the identification as CIvpeaster sp. Several species of CIvpeaster have 
been reported from the Pliocene in Florida, and it is possible the fragments may 
be from disarticulated C. subdepressus , C. crassus , or C. sunnilandensis 
individuals. One slight morphological characteristic preserved in one of the 
fragments is the presence of a small furrow tracing the food groove and 
extending to near the test margin. Such a furrow is present in C. subdepressus . 
and may help in more clearly delineating the species as work continues. A more 
complete fragment or an unbroken individual will be required to interpret with 
certainty the taxonomic identification of these fossils from the Tamiami 
Formation. 


Family Mellitidae Stefanini, 1911 
Genus Encope L. Agassiz, 1840 

Description; Apical system and peristome slightly anterior; posterior 
petals longest: posterior interambulacrum continuous; posterior lunule more than 
half inside line connecting ends of petals. 


166 


Florida species; Up to five taxa are present, including E. aberrans 
Martens, 1867, E. aberrans imperforata Kier, 1963, E. tamiamiensis Mansfield, 
1932, and two possible varieties of Encope sp. from different strata and localities. 

Comments: Fossils of E. aberrans are present in three formations in the 
state, including the Tamiami, Caloosahatchee, and Intracoastal formations. This 
is the first stratigraphic record of the species in the Intracoastal Formation. The 
subspecies E. aberrans imperforata is present in both the Tamiami and 
Caloosahatchee formations. Encope tamiamiensis is present in the Tamiami 
Formation in southern Florida, one of the Encope sp. records is from the 
Tamiami Formation (a new occurrence from Pinecrest Bed unit), the second 
Encope sp. is present in the Nashua Formation, and the final species is present 
in the Jackson Bluff Formation. The fossils from the Nashua Formation are new 
stratigraphic records from the unit, while all three Encope sp. references are 
possible new taxonomic records for the state. Preservation typically is poor for 
all representatives from the three Encope sp. localities, with most specimens 
observed as highly fragmented tests. 

Encope aberrans Martens, 1 867 
(Figure 3-24, A-C and Figure 3-25, A-B) 

Material examined: UF 104520 (figured test), UF 104519 (figured test), 

UF 104518 (figured test), UF 21531 (figured Recent test), UF 104517 (test), UF 
67248 (2 Recent tests). 

Description: Test spade shaped, longer than wide, highest behind the 


apical center, lower surface flat. Typically only two posterior ambulacral notches 


167 


and a lunule, with the three anterior ambulacra only slightly indented. Apical 
system central, star shaped, with five genital pores. Petals broadly lanceolate, 
open; poriferous zones wide, pores conjugate; inner pores circular, larger than 
the outer pores, which tend to enlarge along the conjugations. Peristome small, 
central, circular, with five pairs of buccal tubes. Periproct oval, within the first 
postbasicoronal plates; covered with moveable plates. 

Encope aberrans im perforata Kier, 1 963 
(Figure 3-26, A-B) 

Material examined; UF 56643 (figured test). 

Description; Test broad with width varying from 94 to 101 percent of 
length; test very low varying from 7 to 12 percent of length; greatest width 
posterior to center, anterior margin rounded, posterior sharply truncated; greatest 
height posterior to center; ambulacral notches well developed on some 
specimens, absent on others; posterior closed interambulacral lunule present in 
some specimens thereby preserving area where it would occur, irregularly 
developed, in some specimens opening very small, in others quite large, usually 
irregular in shape, unsymmetrical; in few specimens no lunule; adoral surface 
flat to slightly depressed except for slight elevation between peristome and 
periproct; margin sharp. Apical system slightly anterior, madreporite large, star 
shaped, five genital pores, with genital pore 5 eccentric to right on most 
specimens. Ambulacral petals broad, closing distally, interporiferous zone wider 
in petal III than in other petals; anterior petal III, posterior paired petals (V and I) 
of approximately same length; anterior paired petals shorter than others, in most 


168 


specimens petal II shorter than petal IV. Adoral plate sutures not visible. 
Periproct opening longitudinal, located one-third distance from peristome to 
posterior margin. Peristome located central and circular in shape. This 
subspecies is similar in all respects to the nominate subspecies except that its 
posterior closed lunule is quite small or entirely absent. 

Material examined: UP 8619 (figured test), UP 28217 (140 tests), UP 
29685 (169 tests). 

Description; Plorizontal outline subcircular, concavely truncated behind, 
with four large lateral ambulacral notches and a weaker anterior notch, and a 
rather small posterior lunule; upper surface nearly flat, highest at the front end of 
the lunule; lower surface flat; margin rather thin. Apical system slightly anterior, 
with a large central star-shaped madreporite and five genital pores. The three 
anterior petals lanceolate, extending about two-thirds of the radius, equal; the 
posterior petals longer, curved around the lunule; poriferous zones open, inner 
pores circular, outer pores elongated, pores conjugate. Peristome slightly 
anterior, subcircular. Periproct near the lunule, smaller than the peristome. 
Lunule oval. Pood grooves bifurcating near the peristome, branches slightly 
diverging, nearly straight, obscure lateral branches near the outer ends. Test is 
usually wider than long, rather thin, but not sharp at edges. 

Encope sp. cf. E. aberrans Martens, 1867 
(Pigure 3-27, C-D and Pigure 3-28, A) 

Material examined: UP 104523 (figured test), UP 104524 (figured test in 


matrix). 


169 


Formation: Nashua Formation. 

Locality: Cracker Swamp Ranch 01 (PU004); Putnam County, FL; 
Hastings Quadrangle, SE1/4, SW1/4, Sec. 24, T9S, R27E. 

Collectors: R. Portell and C. Oyen 

Date: 7/9/96 and 7/21/96. 

Description: Test medium size; horizontal outline broadly arcing at 
anterior, widest slightly posterior of center, and truncated at posterior margin in 
interambulacrum 5. Aboral surface gently sloping away from highest point 
posterior of apical system toward margins; margins medium to thin and rounded; 
adoral surface slightly concave from margins toward peristome. Petals large, 
broadly lanceolate, curving nearly closed at distal ends; petals extend 
approximately 85 percent of distance to margin. Apical system at center to 
slightly eccentric anteriorly. Test margin with slight notches at ambulacra II, III, 
and IV; more extensive notches at margins of ambulacra I and IV; all ambulacral 
notches much wider at distal end than at adapical end. Posterior lunule small to 
medium; width proportionally larger thereby producing an elliptical shape. 
Peristome small, circular, located centrally. 

Comments: The fossils collected from this locality most commonly are 
fragmented and show evidence of significant weathering. Several echinoids 
have been discovered in the sediment within bivalves, and they tend to have 
much better preservation and are more complete. Examination of these fossils 
provided the information for the description above, and allows the tentative 
identification of Encope sp. cf. E. aberrans Martens, 1867. This somewhat 


170 


unusual preservation style (echinoids preserved within articulated or isolated 
bivalve shells) is not unique to the Cracker Swamp Ranch locality or the Nashua 
Formation, and similar examples of this preservation have been noted in other 
stratigraphic units (dominantly Neogene strata). In summary, these fossils are 
counted as new stratigraphic records for the Nashua Formation, but are not 
counted as a new taxonomic record or an additional species for the diversity 
summary of the Pliocene. 


cf. Encope sp. 

(Figure 3-28, B-E) 

Material examined: UF 7501 (figured two test fragments), UF 7107 (13 
test fragments), UF 7159 (50 test fragments), UF 79405 (test fragment). 
Formation: Jackson Bluff Formation. 

Locality: Jackson Bluff 02; Leon County, FL; 1 mile west of Bloxham, FL; 
Bloxham Quadrangle, SW 1/4, Sec. 16, T1S, R4W; and Jackson Bluff 01 
(LN001); Leon County, FL; Bloxham Quadrangle, NW 1/4, Sec. 21, T1S, R4W 
Collectors: N. Weisbord (UF 7107, 7159, 7501) and H.K. Brooks (UF 
79405). 

Date: Unknown (Weisbord); 1959 (Brooks). 

Description: Qnly small fragments of test were available for examination. 
Test margin ranges from thinly rounded to broadly rounded, depending on 
fragment examined. Adoral and aboral surfaces relatively flat near margin; some 
evidence for gently increasing slope on aboral surface in petaloid region. 

Petaloid ambulacra potentially broad, lanceolate, and closed at distal end 


171 


(though no complete petals are present in samples). At least one fragment 
shows evidence of an ambulacral notch or lunule, though incomplete. Tubercles 
very small, tightly spaced, and uniformly distributed on plates. No remnants of 
apical system, gonopores, zygopores, peristome, or periproct on fossils collected 
thus far. Test thickness at margin ranges from approximately two to seven mm. 

Comments: These fossils are difficult to identify due the highly 
fragmented nature of the specimens. Most test fragments also are significantly 
weathered deeply enough to expose the stereom, thereby eliminating food 
grooves and most tubercles on the specimens. However, identification to the 
family Mellitidae is reasonable, and based on test margin thickness and potential 
ambulacral invaginations or lunules, I believe these most likely are fragments 
from a species of Encope . Unfortunately, it is not possible to refine the 
identifications until larger, more complete fragments or entire specimens are 
found. These fossils are significant in my study because they are the first 
stratigraphic record of Encope . or any mellitids, from the Jackson Bluff 
Formation. 


Genus Mellita L. Agassiz, 1841 

Description; Thin, flattened, ambitus sharp; paired ambulacral lunules 
only; lunules narrow, elongate, normally closed; anterior paired petals shortest, 
others about equal; peristome and apical system slightly anterior; four genital 
pores; posterior interambulacrum continuous. 


172 


Florida species: M- aclinensis Kier, 1963 and Mellita sp. cf. M- 
caroliniana (Ravenel. 1841), 

Comments; The species M- aclinensis is present in the Tamiami 
Formation, while the Mellita sp. is present in the Nashua Formation. The fossils 
of M. aclinensis are extremely abundant in localized areas and are indicative of 
the Tamiami Formation. The recently collected Mellita sp. cf. M- caroliniana from 
the Nashua represents a new stratigraphic record for the formation, and when 
better preserved specimens are found they may turn out to be new taxonomic 
records too. 


Mellita aclinensis Kier, 1963 
(Figure 3-29, A-B) 

Material examined; UF 28207 (figured test), UF 40359 - 40370 (one test 

each). 

Description: Test margin subcircular except for truncated posterior 
margin on some specimens; width approximately equal to length; test very low 
with thin sharp margin; adoral surface flat to slightly concave; five elongate 
ambulacral lunules in large specimens, lunule in ambulacrum ill smaller than 
others; lunule in posterior interambulacrum very elongate, extending far between 
petals. Apical system slightly anterior, distance from anterior margin to apical 
system approximately 45 percent of length of test; large madreporite; four genital 
pores. Anterior petals II, III, IV lanceolate, straight, petal III longer, extending 
almost two-thirds distance from apical system to anterior margin, petals II and IV 
only halfway to margin; curving posteriorly; in all petals poriferous zone equal in 


173 


width to interporiferous; petals almost closed. Adorally, five pairs of food grooves 
extending from peristome to near margin; area circumscribed by pair of grooves 
expanding distally with greatest width near lunule, constricted distal to lunule; 
area broad between adjacent pairs of grooves. Secondary pores difficult to see 
in most specimens, apparently confined to area circumscribed by food grooves. 
Periproct opening small, elongate, located at anterior edge of lunule. Peristome 
anterior, small, subcircular to pentagonal, food grooves bifurcating near 
peristome. Basicoronal plates small; adoral-most plate of interambulacrum 5 
considerably larger than other basicoronal plates; paired interambulacra 
separated from basicoronal plates by one pair of ambulacral plates, three 
postbasicoronal plates in each column on adoral surface; first pair of 
postbasicoronal interambulacral plates elongate; posterior interambulacrum in 
contact with basicoronal plates; half of periproct within basicoronal 
interambulacrum extending length of lunule. Species characterized by five 
ambulacral lunules. 

Mellita sp. cf. M. caroliniana (Ravenel, 1841) 

(Figure 3-29, C-D) 

Material examined; UF 104526 (figured test in matrix), UF 104527 
(figured test in matrix). 

Formation: Nashua Formation. 

Locality: Cracker Swamp Ranch 01 (PU004); Putnam County, FL; 
Hastings Quadrangle, SE1/4, SW1/4, Sec. 24, T9S, R27E. 

Collectors: R. Portell and C. Oyen. 


174 


Date: 6/25/96. 

Description: Horizontal outline subcircular; widest at approximate 
midpoint to slightly posterior of test. Adoral surface flat; peristome near center; 
aboral surface nearly flat with gentle slope away from apical system. Test 
margin narrowed, varying from pointed to tightly curved. Petals incompletely 
preserved, but lanceolate, nearly closed at distal end; estimated to be 
approximately half the distance to margin, terminating adapically from ambulacral 
lunules; zygopores elliptical, widely spaced, conjugate, with outer pore positioned 
preferentially toward margin. Ambulacral lunules present in each ambulacrum; 
short, oval-shaped; one proportionally larger lunule posterior of periproct in inter- 
mbulacrum 5; narrower than ambulacral lunules. Apical system not preserved in 
specimens. Tubercles small, closely spaced; rarely preserved due to weathering. 

Comments: Weathering and post-mortem transport have left most 
specimens from this locality fragmented and incomplete. The best preserved 
specimens all have been prepared from sediment trapped within large bivalve 
shells (see Figure 3-29, C-D), and these fossils provide the best overall views of 
their morphology. These sand dollars are tentatively identified as Mellita sp. cf. 

M. caroliniana based on the test outline, small ovate ambulacral lunules (five 
total), and the proportionally short petaloid ambulacra. This is the first report of 
M. caroliniana from the Nashua Formation, and I have included these fossils as 
new stratigraphic records for the Pliocene as well as in the total diversity count 
for the epoch. Once specimens are collected which include a preserved apical 


175 

system, peristome, and periproct, comparisons can be completed for all 
morphological features and verification of the identification v\/ill be possible. 

Genus Leodia Gray, 1852 

Description; Like Mellita but with five closed ambulacral lunules. 

Florida species: L. sexiesperforata (Leske, 1778). 

Comments: This species is present in the Nashua Formation. Only two 
fossils have been collected, but they are important specimens. Controversy 
among fellow echinoid workers (zoologists versus paleontologists) leads to a 
tentative identification at this time. I have chosen to assign the fossils to the 
Leodia genus based on the ambulacral lunule arrangement and their overall size. 
The zoologist’s interpretation is that they simply represent unusually large 
individuals of Mellita aclinensis or M. caroliniana . but I find this approach invalid 
based on the fossil M- aclinensis individuals I have sampled and measured. 

None of the hundreds of M- aclinensis fossils from Florida are close to the size of 
these specimens from the Nashua Formation, and therefore I believe the 
difference is real rather than a simple artifact. Continued collection and sampling 
from the Nashua may provide new specimens in the future, which can help solve 
the taxonomic problem associated with these fossils. 

Leodia sexiesperforata (Leske, 1778) 

(Figure 3-30, A) 

Material examined: UF 31969 (figured test). 


176 

Description: Horizontal outline subcircular, flattened behind; upper 
surface nearly flat, highest at the anterior petal; lower surface flat. Apical system 
central; madreporite star shaped, with five sharp points; four genital pores 
outside the madreporite and detached from it. Petals short, less than half the 
radius, nearly equal in length, lanceolate; poriferous zones as wide as the 
interporiferous, inner pores oval, outer pores elongated, pores conjugate. All five 
ambulacra penetrated by a long, narrow, straight lunule between the petals and 
the margin; posterior lunule slightly wider, occupying the middle third of the 
radius. Peristome small, circular, central, with paired buccal tubes. Periproct 
small, pear-shaped, lying midway between the peristome and the lunule. Food 
grooves deep, narrow, perforated; divaricating at the peristome, each branch 
running parallel to that of the neighboring ambulacrum and branching several 
times near the margin. Spines short, straight. 

Order Cassiduloida Claus, 1880 
Family Cassidulidae L. Agassiz and Desor, 1847 
Genus Rhvncholampas A. Agassiz, 1869 

Description: See Oligocene echinoid section for generic description. 

Florida species: Two species are found in the state, including R. avresi 
Kier, 1963 and R. everqiadensis (Mansfield, 1932). 

Comments: The species R. everqiadensis is present in the Tamiami 
Formation and R. avresi is present in the Caloosahatchee Formation. 


Rhyncholampas ayresi Kier, 1963 
(Figure 3-30, B-C) 


177 


Material examined; UF 63062 (figured test), UF 63215 (test), UF 63368 
(test), UF 63767 (test), UF 63770 (test). 

Description: Test yarying in length; test width approximately 85 to 90 
percent of length with greatest width posterior of center; adapical surface highly 
inflated with steeply sloping sides, height ayeraging 55 percent of length; adoral 
surface flat or slightly depressed around peristome. Apical system anterior, four 
genital pores, compact. Petals well deyeloped, broad, lanceolate, with greatest 
width one-third distance from apical system to end of petal, all petals of 
approximately equal length, petals II, IV wider than others, petal III narrower; 
poriferous zones of unequal length with one to three more pore pairs in right 
poriferous zone of petal II, posterior zones of petals II and IV, and anterior 
poriferous zones of petals V and I; single pores in ambulacral plates beyond 
petals. Periproct supramarginal, wider than high, with slight grooye extending 
from opening to posterior margin. Peristome anterior, pentagonal, depressed, 
wider than high. Floscelle with phyllodes well deyeloped, broad, approximately 
30 pores in each phyllode, with 10 in each outer series, four to six irregularly 
arranged in each inner. Buccal pores present. Bourrelets yery prominent, 
pointed. Tubercles adorally much larger than adapically, narrow naked granular 
zone in median area of interambulacrum 5 and ambulacrum III adorally. Species 
characterized by highly inflated adapical surface, steep sides, smooth marginal 
outline, narrow naked zone in interambulacrum 5, and narrow phyllode III. 


178 

Rhyncholampas everqiadensis (Mansfield. 1932) 

(Figure 3-31, A-B) 

Material examined; UF 17312 (figured test), UF 20918 - 20925 (one test 
each lot), UF 34405 (test), UF 60126 (test). 

Description; Test large, suborbicular, and moderately high; upper 
surface convex and broadly rounded, the posterior surface more gently inclined 
than the anterior; lower surface nearly flat except in the area surrounding the 
peristome, where it is shallowly concave. Apical system, situated opposite the 
peristome, is rather large, granular, and slightly elevated; a genital pore is at the 
juncture of the petals and a smaller radial pore is opposite each petal. 

Ambulacral areas petaloid at dorsal portions. Petals rather long, extending 
nearly to the ambitus, expanding to about one-third their length from the apical 
system, then gradually contracting distally, and nearly closing at their extremities; 
poriferous zones rather wide, shallowly depressed; pores nearly equal in size 
and rounded in outline; pairs of pores conjugate. Interporiferous areas weakly 
tumid. Posterior interambulacrum weakly medially arched. Periproct rather 
large, longest transversely; supramarginal, the lower margin being about four 
millimeters above the ambitus; the upper arched margin slightly overhangs the 
aperture. Peristome eccentric anteriorly, pentagonal, transversely elongate, and 
surrounded by a large well-defined floscelle with prominent bourrelets. The outer 
pores of the floscelle are more direct and more regularly placed; the inner ones 
are more irregularly placed and some of them are arranged in two rows. The 
surface of the test is closely set with scrobiculate tubercles. 


179 

Order Spatangoida Claus, 1876 
Family Paleopneustidae A. Agassiz, 1904 
Genus Pericosmus L. Agassiz, 1847 

Description: Peripetalous fascicle passing above periproct and entirely 
separate marginal fascicle passing below periproct, peripetalous fascicle may 
branch anteriorly, and one or other fascicle may disappear anteriorly; apical 
system ethmolytic, with three or four gonopores; paired ambulacra having 
depressed petals which tend to have distal plates occluded. 

Comments: One or possibly two species were collected from an 
undetermined stratigraphic unit in the Gulf of Mexico, and herein are referred to 
as Pericosmus spp. 


Pericosmus spp. 

(Figure 3-31, C-F and Figure 3-32, A-B) 

Material examined: UF 101885 (figured partial, phosphatized test with 
nearly complete internal mold), UF 66566 (figured internal mold of test). 

Formation: Uncertain. 

Locality: Ocean floor samples. Gulf of Mexico. UF 101885 dredged from 
approximately 511 m, December 19, 1989 at OTB5 (27°01’N, 84° 56’W) (UF 
locality 3784). The second fossil, UF 66566, dredged from approximately 520 m. 
May 4, 1993, at WFS1 (26° 56.29’N, 84° 55.75’W) (UF locality 3810). 

Collectors: D. Hodell (UF 101885) and K. Fountain (UF 66566). 

Date; 12/19/89 (UF 101885) and 5/4/93 (UF 66566). 

Description: One fossil, UF 101885, test outline is subcircular, with 
maximum width slightly anterior of ambitus. Specimen somewhat compressed at 


180 


apical system, although overall shape is not distorted significantly. Unfortunately, 
this compaction and the remnant sediment cemented to the surface has inhibited 
identification of apical system morphology. Periproct and peristome are nearly 
intact and largely unaffected by diagenesis or fragmentation. Peristome has 
elevated labrum, and is located slightly anterior of the test center and distinctly 
posterior of the anterior sulcus termination on adoral surface. Anterior sulcus is 
modestly developed and approximately half as deep as it is v\/ide. Adoral surface 
is generally flat while aboral surface originally was dome shaped (though 
presently collapsed). Test margins are broadly rounded. The anterior half of 
both adoral and aboral surfaces show well-preserved molds of tubercles, both 
large and small, and relict plate sutures are visible in various areas of the 
specimen. Ambulacrum I and IV petals are partially preserved, with pore-pairs 
present. Petals are closed, lanceolate, and terminate approximately midway 
between former apical system and ambitus. The second fossil, UP 66566, test 
outline subcircular to subpentagonal, with maximum test width located distinctly 
anterior of ambitus. All surfaces of this internal mold have been bored 
extensively, but the general shape of the test is preserved. Adoral surface 
relatively flat, with test margins broadly curved or rounded, and a gently domed 
aboral surface with a slight peak at the approximate apical system location. No 
ambulacra, plate sutures, tubercles, or other morphological traits are visible on 
the mold surfaces. Anterior sulcus is small to moderate in size, with sulcus 
length subequal in dimension to the sulcus width. Peristome visible on adoral 
surface, positioned distinctly anterior of test center and near the posterior margin 


181 


of sulcus. Peristome slightly curved and ovate, with elevated labrum. Periproct 
not preserved. 

Comments: The UP 101885 specimen has undergone phosphatization, 
but it is apparent that the anterior portion of the test is intact while only the 
posterior portion is exclusively an internal mold of phosphatized carbonate 
sediment. Some micromorphology is visible, such as tubercles and pore-pairs 
(see description), and overall preservation is the best of the echinoids collected 
along the upper west Florida slope. Surface borings and epibionts are present, 
and are located principally on the aboral surface. The only significant 
preservationai effect which limits complete description of the specimen is the 
slight compaction and collapse of the apical system. 

Preservation quality of the second Pericosmus specimen (UF 66566) is 
better than that of the brissid (UF 57743; see below), but it is a more poorly 
preserved internal mold than the other Pericosmus sp. fossil (UF 101885); 
therefore, relatively few diagnostic morphological characteristics could be 
described. However, in addition to the traits of test length, width, and height, the 
shape and position of the peristome are evident. The adoral surface of UF 
66566 (Figure 8A) is imperfectly preserved and bored, but adorally (Figure 8C) 
the mold provides a generally good surface for examination. Since the peristome 
shape and position commonly are used to aid in taxonomic descriptions, this 
specimen can be identified (tentatively) to generic level. Based on these 
discernable characteristics, the best placement for this specimen is also within 
the genus Pericosmus . Oyen et al. (2000) discuss these fossils in more detail. 


182 


The primary significance of these fossils is a function of their stratigraphic 
age and geographic distribution. Both fossil and extant species of Pericosmus 
are known from the Caribbean as well as from other parts of the world. Only 
three countries in the Caribbean and the Gulf of Mexico region, Cuba, Costa 
Rica, and Venezuela, have fossil records of this genus. All of the Cuban species 
were found in Eocene through Miocene age strata. The Venezuelan and Costa 
Rican species both were collected from the Miocene. Therefore, these 
specimens represent the youngest fossils of Pericosmus in the Caribbean and 
Gulf of Mexico. In addition, only one other record of a Pliocene species (i.e., P. 
schencki Israelsky, 1933 from the Malumbang Formation, Philippine Islands) has 
been published. Because the age of the Philippine fossil is very questionable, 
these upper west Florida slope fossils may provide a biostratigraphic range 
extension of the genus. Furthermore, if my identification is correct, this 
represents the first report of fossil Pericosmus from the United States. 

Family Schizasteridae Lambert, 1905 
Genus Aqassizia L. Agassiz and Desor, 1847 

Description: See Eocene echinoid section for generic description. 

Florida species; A. porifera (Ravenel, 1848). 

Comments; The species is present in the Caloosahatchee and Tamiami 
formations. The Tamiami Formation occurrence reported herein is a new 
stratigraphic record of the species that was collected from a pit in southwestern 


Florida. 


Aqassizia porifera (Ravenel, 1848) 
(Figure 3-33, A-B) 


183 


Material examined: UF 12894 (figured test), UF 14144 (test), UF 22150 
(test), UF 23973 (test), UF 24522 (test). 

Description: Florizontal outline ovate, widest in front, truncated behind; 
upper surface strongly inflated to subconical, highest at the apical system, 
sloping steeply forward, less steeply to the posterior truncation; margin broadly 
rounded; lower surface gently inflated. Apical system nearly central; four genital 
pores, rather close together; ethmolytic, the madreporite extending beyond the 
posterior ocular pores. Anterior ambulacrum very slightly depressed, not at all 
depressed at the margin; pores obscure. Paired petals moderately depressed; 
long, the posterior pair somewhat shorter; anterior paired petals in line near the 
apex, curving gently forward to an angle of approximately 97°; posterior pair 
curving slightly outward to an angle of approximately 70°; pores of posterior 
petals and posterior zone of anterior pair very small and inconspicuous, pore 
pairs oblique; interporiferous zones narrow. Peristome at the anterior quarter, 
reniform; labium large. Periproct large, transversely elliptical; at the top of 
truncation, which is depressed. Plastron expanding for more than half its length, 
then sides curving inward. Marginal fascicle complete, curving downward below 
the periproct; hemipetalous fascicle indented in the posterior and postero-lateral 
interambulacra, meeting the marginal fascicle behind the anterior paired petals. 

Family Brissidae Gray, 1855 
Genus Plagiobrissus Pomel, 1883 


Description: See Eocene echinoid section for generic description. 


Florida species: P. qrandis (Gmelin, 1791). 

Comments: This species is present in the Tamiami Formation. It is a 


184 


unique specimen that may represent the first fossil record of this species, and 
therefore it represents a new stratigraphic record for the species. Caution must 
be exercised with this fossil, however, due to the incomplete nature of the test. 

Plaqiobrissus qrandis (Gmelin. 1791) 

(Figure 3-33, C-E) 

Material examined: UF 22152 (figured incomplete test), UF 5343 
(Recent test). 

Description: Test very large; horizontal outline suboval, truncated and 
somewhat emarginate in front; upper surface flat on top, steeply sloping at each 
end; lower surface gently convex; margin acutely rounded. Apical system slightly 
anterior; proportionately small; four genital pores; strongly ethmolytic, the 
madreporite extending far beyond the ocular plates. Anterior ambulacrum 
narrow, depressed; plates nearly equilateral; pore pairs small, longitudinal. 

Petals narrow, somewhat flexuous, depressed, extending more than two-thirds 
the way to the margin; anterior pair diverging at an angle of 100°, posterior pair, 
37°; pores circular, strongly conjugate; interporiferous zones narrower. 

Ambulacra on lower surface very narrow; posterior pair nearly parallel. 

Peristome at the anterior quarter, semilunate; floscelle conspicuous. Paired 
interambulacra excluded from the peristome. Periproct submarginal, sloping 
upward and backward, not visible from above; longer than wide. Escutcheon 
semilunate, concave behind; surrounded by a wide subanal fasciole. Anal 


185 


fasciole broadly U-shaped, wider than high, branches extending about even with 
the upper end of the periproct. Peripetalous fasciole narrow, nearly oval. 
Tubercles of paired interambulacra large, perforated; confined within the 
peripetalous fasciole. Tubercles of posterior interambulacrum somewhat smaller, 
confined to median region but extending beyond the fasciole almost to the 
posterior end. 


Family Brissidae Gray, 1855 
Fam., gen., et sp. indet. 

(Figure 3-34 A-C) 

Material examined; UF 57743 (figured internal mold of test). 

Formation; Uncertain. 

Locality; Ocean floor sample. Gulf of Mexico. Dredged from 
approximately 511 m, at OTB8 (27° 05’N, 84° 57’W) (UF locality 3811). 

Collector; K. Fountain. 

Date; 12/19/89. 

Description; Very limited morphological features are visible on this fossil, 
although the general test length, width, and height can be distinguished, as well 
as the slight sulcus along ambulacrum III. 

Comments; Of the three echinoids recovered from the upper west Florida 
slope, this specimen is the most poorly preserved. The relative proportions of 
the test length, width, and height, the slight anterior sulcus, and the overall shape 
suggest a strong affinity to the Brissidae (see Oyen et al., 2000 for further 


discussions). 


186 

Family Loveniidae Lambert, 1905 
Genus Echinocardium Gray, 1825 

Description: Differs from typical loveniids in scarcity of large spines and 
tubercles, and absence of deep areoles or camellae; subanal fasciole with pair of 
anal branches. 

Florida species: E. orthonotum (Conrad, 1843). 

Comments: The species is present in the Tamiami, Jackson Bluff, and 
Intracoastal formations. This stratigraphic distribution includes a new 
stratigraphic record from the Intracoastal Formation of northern Florida. Readers 
should note that Kier (1963) originally reported this fossil in Florida as E. 
qothicum (Ravenel, 1848), which was an incorrect specific assignment on Kier’s 
part. Also, the preservation of the Jackson Bluff specimen is poor, and my 
identification of the species should be considered only a preliminary 
identification. 


Echinocardium orthonotum (Conrad, 1843) 

(Figure 3-34, D-H) 

Material examined: UF 60182 (figured test), UF 104522 (figured test), 

UF 62736 (5 test fragments), UF 62739 (test), UF 39540 (16 test fragments). 

Description: Test ovate, convex-depressed; truncated at each end, more 
elevated anteriorly than posteriorly; dorsal line of the suture a little elevated, and 
curved gradually to the mouth on the anterior half; on the posterior, straight to the 
margin and parallel to the base; canal very wide and slightly impressed on the 
back, margined by an obtuse carinated line and slight furrow; on the periphery 


Figure 3-19. Pliocene regular echinoids. 

A) Eucidaris tribuloides (Lamarck, 1816); UF 72022; aboral view of flattened 
test; Tamiami Formation; lx. 

B) Eucidaris tribuloides (Lamarck. 1816); UF 72022; adoral view of flattened test 
showing partially exposed lantern; Tamiami Formation; lx. 

C) Arbacia imorocera (Conrad. 1843); UF 83420; aboral view of test; Tamiami 
Formation; lx. 

D) Arbacia sp.; UF 7506; aboral view of test; Jackson Bluff Formation; lx. 

E) Arbacia sp.; UF 7506; adoral view of test; Jackson Bluff Formation; lx. 

F) Arbacia sp. cf. A. imorocera (Conrad, 1843); UF 7507; aboral view of test 
fragment; Jackson Bluff Formation; lx. 

G) cf. Arbacia sp.; UF 84283; aboral view of test fragment; Nashua Formation; 
lx. 

FI) Lvtechinus varieqatus olurituberculatus Kier, 1 963; UF 1 2895; aboral view of 
test; Caloosahatchee Formation; lx. 

I) Lvtechinus varieqatus olurituberculatus Kier, 1963; UF 12895; adoral view of 
test; Caloosahatchee Formation; lx. 

J) Echinometra lucunter (Linnaeus, 1758); UF 12937; aboral view of test; 
Caloosahatchee Formation; lx. 

K) Echinometra lucunter (Linnaeus, 1758); UF 12937; adoral view of test; 
Caloosahatchee Formation; lx. 


188 



Figure 3-20. Pliocene irregular echinoids. 

A) CIvpeaster sunnilandensis Kier, 1963; UF 22148; aboral view of test; 
Tamiami Formation; 0.75x 

B) CIvpeaster sunnilandensis Kier, 1963; UF 22148; adoral view of test; 
Tamiami Formation; 0.75x 


190 



Figure 3-21. Pliocene irregular echinoids. 

A) CIvpeaster rosaceus dalli (Twitchell, 1915); UF 65813; aboral view of test; 
Caloosahatchee Formation; 0.75x. 

B) CIvpeaster rosaceus dalli (Twitchell, 1915); UF 65813; adoral view of test; 
Caloosahatchee Formation; 0.75x. 


192 



Figure 3-22. Pliocene irregular echinoids with Recent specimens for comparison. 

A) Clvoeaster subdeoressus (Gray. 1825); UF 98692; aboral view of test; 
Tamiami Formation; 0.75x. 

B) Clvoeaster subdeoressus (Gray, 1 825); UF 21532, aboral view of Recent 
test; 0.5x. 

C) Clvoeaster subdeoressus (Gray. 1825); UF 21532, adoral view of Recent 
test; 0.5x. 


194 



Figure 3-23. Pliocene irregular echinoids. 

A) CIvpeaster sp.; UF 104521; adoral view of incomplete test; Intracoastal 
Formation; lx. 

B) CIvpeaster sp.; UF 30870; aboral view of test fragment; Tamiami Formation; 
lx. 

C) CIvpeaster sp.; UF 30870; adoral view of test fragment; Tamiami Formation; 
lx. 

D) CIvpeaster sp.; UF 44202; aboral view of test fragment; Tamiami Formation; 
lx. 

E) CIvpeaster sp.; UF 44202; adoral view of test fragment; Tamiami Formation; 
lx. 


196 



Figure 3-24. Pliocene irregular echinoids. 

A) Encope aberrans Martens. 1867; UF 104520, aboral view of test fragment; 
Intracoastal Formation; 0.57x. 

B) Encope aberrans Martens. 1867; UF 104519, adoral view of test; Intracoastal 
Formation; 0.75x. 

C) Encope aberrans Martens, 1867; UF 104518, adoral view of test; Intracoastal 
Formation; 0.75x. 


198 





Figure 3-25. Recent irregular echinoids for comparison purposes. 

A) Encope aberrans Martens. 1867; UF 21531; aboral view of Recent test; lx 

B) Encope aberrans Martens, 1867; UF 21531; adoral view of Recent test; lx 


200 




Figure 3-26. Pliocene irregular echinoids. 

A) Encope aberrans imperforata Kier, 1 963; UF 56643; aboral view of highly 
eroded test; Caloosahatchee Formation; 1x. 

B) Encope aberrans imperforata Kier, 1963; UF 56643; adoral view of highly 
eroded test; Caloosahatchee Formation; lx. 


202 



Figure 3-27. Pliocene irregular echinoids. 

A) Encope tamiamiensis Mansfield. 1932; UF 8619; aboral view of test; Tamiami 
Formation; lx. 

B) Encope tamiamiensis Mansfield, 1 932; UF 8619; adoral view of test; Tamiami 
Formation; lx. 

C) Encope sp. cf. E. aberrans Martens, 1867; UF 104523; aboral view of test; 
Nashua Formation; lx. 

D) Encope sp. cf. E. aberrans Martens, 1867; UF 104523; adoral view of test; 
Nashua Formation; lx. 


204 



Figure 3-28. Pliocene irregular echinoids. 

A) Encope sp. cf. E. aberrans Martens, 1867; UF 104524; aboral view of test 
Nashua Formation; lx. 

B) cf. Encope sp.; UF 7501 ; test fragment; Jackson Bluff Formation; lx. 

C) cf. Encope sp.; UF 7501 ; test fragment; Jackson Bluff Formation; lx. 

D) cf. Encope sp.; UF 7501 ; test fragment; Jackson Bluff Formation; lx. 

E) cf. Encope sp.; UF 7501 ; test fragment; Jackson Bluff Formation; lx. 


206 




Figure 3-29. Pliocene irregular echinoids. 

A) Mellita aclinensis Kier, 1 963; UF 28207; aboral view of test; Tamiami 
Formation; 1x. 

B) Mellita aclinensis Kier, 1963; UF 28207; adoral view of test; Nashua 
Formation; lx. 

C) Mellita sp. of. M- caroliniana (Ravenel, 1841); UF 104526, adoral view of test; 
Nashua Formation; 0.63x; 

D) Mellita sp. of. M- caroliniana (Ravenel, 1841); UF 104527, adoral view of test; 
Nashua Formation; 0.65x 


208 



Figure 3-30. Pliocene irregular echinoids. 

A) Leodia sexiesoerforata (Leske, 1778); UF 31969; aboral view of test; 
Tamiami Formation; 0.75x 

B) Rhvncholampas avresi Kier. 1963; UF 63062; aboral view of test; 
Caloosahatchee Formation; lx. 

C) Rhvncholampas avresi Kier, 1963; UF 63062; adoral view of test; 
Caloosahatchee Formation; lx. 


210 



Figure 3-31 . Pliocene irregular echinoids. 

A) Rhvncholampas everqiadensis (Mansfield, 1932); UF 17312; aboral view of 
test; Tamiami Formation; lx. 

B) Rhvncholampas everqiadensis (Mansfield, 1932); UF 17312; adoral view of 
test; Tamiami Formation; lx. 

C) Pericosmus sp.; UF 101885; aboral view of phosphatized, partial test and 
internal mold; undetermined stratigraphic unit; lx. 

D) Pericosmus sp.; UF 101885; adoral view of phosphatized, partial test and 
internal mold; undetermined stratigraphic unit; lx. 

E) Pericosmus sp.; UF 101885; anterior view of phosphatized, partial test and 
internal mold with peristome visible; undetermined stratigraphic unit; lx. 

F) Pericosmus sp.; UF 101885; posterior view of phosphatized, partial test and 
internal mold with outline of periproct visible; undetermined stratigraphic unit; 
lx. 


212 



Figure 3-32. Pliocene irregular echinoids. 

A) Pericosmus sp.; UF 66566; aboral view of phosphatized, partial internal mold 
of test; undetermined stratigraphic unit; 1x. 

B) Pericosmus sp.; UF 66566; adoral view of phosphatized, partial internal mold 
of test; undetermined stratigraphic unit; 1x. 


214 



Figure 3-33. Pliocene irregular echinoids. 

A) Aqassizia porifera (Ravenel, 1848); UF 12894; aboral view of test; 
Caloosahatchee Formation; lx. 

B) Aqassizia porifera (Ravenel, 1848); UF 12894; adoral view of test; 
Caloosahatchee Formation; lx. 

C) Plaqiobrissus qrandis (Gmelin, 1791); UF 22152; aboral view of partial test; 
Tamiami Formation; 0.75x. 

D) Plaqiobrissus qrandis (Gmelin, 1791); UF 22152; adoral view of partial test; 
Tamiami Formation; 0.75x. 

E) Plaqiobrissus qrandis (Gmelin. 1791); UF 22152; posterior view of test with 
the outline of periproct visible; Tamiami Formation; 0.75x. 


216 



Figure 3-34. Pliocene irregular echinoids. 

A) BRISSIDAE; UF 57743; aboral view of phosphatized, internal mold of test; 
undetermined stratigraphic unit; 1x. 

B) BRISSIDAE; UF 57743; adoral view of phosphatized, internal mold of test; 
undetermined stratigraphic unit; 1x. 

C) BRISSIDAE; UF 57743; lateral view of phosphatized, internal mold of test; 
undetermined stratigraphic unit; 1x. 

D) Echinocardium orthonotum (Conrad, 1 843); UF 601 82; aboral view of test; 
Tamiami Formation; lx. 

E) Echinocardium orthonotum (Conrad, 1 843); UF 60182; adoral view of test; 
Tamiami Formation; lx. 

F) Echinocardium orthonotum (Conrad, 1843); UF 104522; aboral view of test; 
Intracoastal Formation; lx. 

G) Echinocardium orthonotum (Conrad. 1843); UF 104522; adoral view of test; 
Intracoastal Formation; lx. 

H) Echinocardium orthonotum (Conrad. 1843); UF 104522; posterior view of test 
with outline of periproct visible; Intracoastal Formation; lx. 

I) Echinocardium sp. cf. E. orthonotum (Conrad, 1843); UF 84281; aboral view of 
exterior of test fragment; Nashua Formation; lx. 

J) Echinocardium sp. cf. E. orthonotum (Conrad, 1843); UF 84281; aboral view 
of interior of test fragment; Nashua Formation; lx. 


218 




219 


the canal is deep and angular; ambulacra rapidly expanding from the extremities 
towards the dorsal suture; pores disunited; in the middle of the back a slight 
furrow crosses obliquely each of the anterior ambulacra at its termination; base 
plano-convex; anus large and remote from the margin; granulation on the back 
minute and very closely arranged, in the canal much smaller and more closely 
arranged towards the margins. 

Echinocardium sp. cf. E. orthonotum (Conrad, 1843) 

(Figure 3-34, l-J) 

Material examined: UF 84281 (figured test fragment). 

Formation: Nashua Formation. 

Locality: Cracker Swamp Ranch 01 (PU004); Putnam County, FL; 
Hastings Quadrangle, SE1/4, SW1/4, Sec. 24, T9S, R27E. 

Collectors: R. Portell, C. Oyen, et al. 

Date: 6/26/96. 

Description: One small test fragment, dominantly of an ambulacral 
region (either ambulacrum I or II). Test plates thin; tubercles small, densely 
spaced throughout. Distal end of ambulacrum with pore pairs curving slightly 
anterior. Pore pairs circular, large, widely separated and conjugate; two to three 
pores per ambulacral plate; petaloid structure moderately depressed. Ambitus 
broadly curving toward adoral surface (surface not present in sample, however). 

Comments: Although very little of the original test was collected, the 
portion described matches well with the genus Echinocardium (see generic 
description provided earlier in Pliocene section). The fossil displays a similar 


220 


ambulacral structure as that of E, orthonotum . which is found in other Pliocene 
strata in Florida. Yet, just as in other cases of fragmented fossil echinoids, this 
identification must remain tentative until more complete specimens are available 
for comparison. Therefore, herein I refer to this specimen as Echinocardium sp. 
cf. E. orthonotum and include it as a new stratigraphic record for the Nashua 
Formation (both at the generic and specific levels) but do not include it as a new 
taxonomic record or as an additional species for the Pliocene diversity count. 

Pleistocene Echinoids 

Order Clypeasteroida A. Agassiz, 1872 
Family Clypeasteridae L. Agassiz, 1835 
Genus Clypeaster Lamarck, 1801 

Description: See Oligocene echinoid section for generic description. 

Florida species: Up to three species are present, including C. rosaceus 
(Linnaeus, 1758), C. rosaceus dalli (Twitchell, 1915), and an unidentified 
Clypeaster sp. 

Comments: All fossils of Clypeaster are present in the Bermont 
Formation. The unidentified fossil (referred to as Clypeaster sp.) appears to be a 
new species, and therefore represents both a new stratigraphic and taxonomic 
record for the state. 


Clypeaster rosaceus (Linnaeus, 1758) 

(Figure 3-35, A-B) 

Material examined: UF 42000 (figured test), UF 70612 (test), UF 63527 
(test), UF 63940 (test), UF 63949 (test). 


221 


Description: Test large; horizontal outline subpentagonal; upper surface 
strongly inflated, with swollen petals; lower surface flat near the margin, with five 
conspicuous ambulacral grooves, deeply concave around the peristome; margin 
broadly rounded. Apical system central, with a large central madreporite; genital 
pores five, usually outside the apical system. Petals long, broad, swollen, 
completely closed at the apex, moderately open distally; poriferous zones 
proportionately narrow, strongly curved inward near the tips; pores circular, 
conjugate. Peristome central, deeply sunken, pentagonal. Periproct small, 
circular, submarginal. Tubercles sunken in small scrobicules, perforated. 

Clypeaster rosaceus dalli (Twitchell, 1915) 

(see Figure 3-21, A-B in Pliocene section) 

Material examined: UF 63222 (test), UF 63300 (2 tests). 

Description: See specific description in Pliocene section. 


Clypeaster sp. 

(Figure 3-35, C-D) 

Material examined: UF 54188 (figured test). 

Formation: Bermont Formation. 

Locality: South Bay 04 (PB007), USA, Florida, Palm Beach County, 
Everglades 1 NW/ Everglades 1 NE Quadrangles, T46S, R37E. 

Collector: McGinty Collection. 


Date: 3/31/1968. 


222 


Description: Test small to medium size; horizontal outline subovate to 
subpentagonal with minimal marginal notches at interambulacra 1 and 4 and 
margin depressed slightly at interambulacrum 5. Anterior margin thick; posterior 
margin somewhat thinner. Aboral surface tumid, moderately raised in petaloidal 
region and apical system; margin broadly rounded; adoral surface moderately 
concave, increasing near peristome, with paired ambulacral furrows distinct and 
curving slightly toward each other at peristome. Apical system slightly broken, 
but located centrally, above peristome. Petals relatively short, extending about 
50% of distance to margin; subequal length, with petaloid ambulacrum III only 
slightly longer; pore pairs close distinctly adapically, essentially disappearing, 
broaden two-thirds of way to end, then close at ends; pores conjugate; 
interporiferous zones broad. Peristome central, circular. Periproct inframarginal, 
small, round. Tubercles small. 

Comments: This fossil is very well preserved and, after careful 
examination, compares most closely with C. subdepressus . However, it differs in 
overall size by being much smaller than adult C. subdepressus individuals, and 
secondly, the adoral surface is significantly more concave than the typical C. 
subdepressus . Unfortunately, only one specimen has been collected and until 
more collecting can be done in this unit to obtain additional specimens, I have 
chosen to leave this specimen identified only as Clypeaster sp. I believe the 
morphological differences warrant including this as a potentially new taxonomic 
record, as well as a new stratigraphic record (regardless of whether it is a new 
species or simply an unusual specimen of C. subdepressus) . 


223 


Family Mellitidae Stefanini, 1911 
Genus Encope L. Agassiz, 1840 

Description: See Pliocene echinoid section for generic description. 

Florida species: Two species are present, including E. aberrans 
Martens, 1867 and E. michelini L. Agassiz, 1841. 

Comments: Both species are present in the Bermont Formation, and E. 
michelini also is present in the Anastasia Formation. The report herein of E. 
michelini in the Anastasia is a new stratigraphic record of this species. Both 
species of echinoids typically are fragmented, with specimens of E. michelini 
infrequently preserved as intact, whole fossils (excluding spines). 

Encope aberrans Martens, 1867 
(Figure 3-36, A-B) 

Material examined: UF 42001 (figured test). 

Description: Test spade shaped, longer than wide, highest behind the 
apical center, lower surface flat. Typically only two posterior ambulacral notches 
and a lunule, with the three anterior ambulacra only slightly indented. Apical 
system central, star shaped, with five genital pores. Petals broadly lanceolate, 
open; poriferous zones wide, pores conjugate; inner pores circular, larger than 
the outer pores, which tend to enlarge along the conjugations. Peristome small, 
central, circular, with five pairs of buccal tubes. Periproct oval, within the first 
postbasicoronal plates; covered with moveable plates. 


224 


Encope michelini L. Agassiz, 1 841 
(Figure 3-36, C-D and Figure 3-37, A-B) 

Material examined; UF 62981 (figured test), UF 101080 (figured test). 


UF 64984 (test), UF 64985 (test), UF 67082 (3 tests), UF 67217 (2 tests). 

Description: Florizontal outline elliptical, truncated behind; upper surface 
gently tumid, usually higher in front than behind; lower surface flat; margin thin. 
Posterior lunule long and rather wide. Five deep ambulacral notches, which tend 
to become oval and to close at the outer end. Apical system anterior; 
madreporite star shaped; five genital pores. Petals broadly lanceolate, extending 
more than halfway to the margin; poriferous zones wide, curved together at the 
outer ends but not closed. Peristome below the apical system, circular, with five 
pairs of buccal tubes. Food grooves diverging near the peristome and curving 
together around the notches; several branches near the margin. Periproct near 
the lunule. 


Genus Mellita L. Agassiz, 1841 

Description: See Pliocene echinoid section for generic description. 
Florida species: M quinquiesperforata (Leske. 1778). 

Comments: The species is present in two Pleistocene formations; the 
Anastasia Formation and the Satilla Formation. Specimens typically are poorly 
preserved and often fragmented, though specimens have been collected intact 
less frequently. This species represents a new stratigraphic record from the 


Satilla Formation of Florida. 


Mellita quinquiesperforata (Leske, 1778) 

(Figure 3-38, A-B) 

Material examined: UF 14778 (figured test), UF 84156 - 84221 (one 


225 


Recent test for each UF number). 

Description: Fiorizontal outline subcircular, usually flattened behind and 
weakly notched in front; upper surface nearly flat, sloping evenly in all directions 
from the apical system to the very thin margin; lower surface flat. Test perforated 
by narrow radial slots near the outer ends of the paired ambulacra and by a 
longer slot in the median part of the posterior interambulacrum. Apical system 
having five small ocular pores; four genital pores at the paired interambulacral 
tips of the large star-shaped madreporite. Petals extending about halfway to the 
margin, anterior paired petals slightly shorter and more rounded than the others; 
poriferous zones about as wide as the interporiferous zones, open at the rounded 
tips. Peristome small, central. Periproct elongated, midway between the 
peristome and the posterior slot. Food grooves narrow, shallow, divaricating 
near the peristome, the branches nearly surrounding the ambulacra, starting from 
five nodes, each covering twin buccal tubes. Surface covered with short acicular 
spines, which are longest around the perforations and on the lower surface. 

Order Spatangoida Claus, 1876 
Family Schizasteridae Lambert, 1905 
Genus Moira A. Agassiz, 1872 

Description: Distinguished from Schizaster by deeply sunken nature of it 

petals. 

Florida species: M. atropos (Lamarck, 1816). 


226 

Comments; The species is present in the Bermont Formation. This 
represents a new stratigraphic record of the species in Florida. In most instances 
fossils are highly fragmented, which probably has resulted in the lack of pub- 
lished information about this species’ presence in the fossil record. 

Moira atropos (Lamarck. 1816) 

(Figure 3-39, A-H) 

Material examined: UF 100180 (figured test), UF 12641 (figured Recent 

test). 

Description: Horizontal outline suboval with an anterior notch, truncated 
behind; upper surface swollen behind, sloping steeply forward from the apical 
system; margin rounded; lower surface rounded. Apical system slightly posterior; 
two genital pores far apart, behind the anterior paired ocular pores but in line with 
the anterior ocular pore. Petals deeply sunken, depressions constricted at the 
top; posterior paired petals much shorter than the anterior pair. Peristome at the 
anterior quarter, reniform, strongly lipped. Periproct high on the posterior 
truncation, longer than wide. Peripetalous fasciole deeply indented in all five 
interambulacra, lying near the edges of the petals; lateral fascioles normal. 

Sutures bare. Long, curved spines. 


Figure 3-35. Pleistocene irregular echinoids. 

A) CIvpeaster rosaceus (Linnaeus, 1 758); UF 42000; aboral view of test; 
Bermont Formation; 0.75x. 

B) CIvpeaster rosaceus (Linnaeus, 1758); UF 42000; adoral view of test; 
Bermont Formation; 0.75x. 

C) CIvpeaster sp.; UF 54188; aboral view of test; Bermont Formation; lx. 

D) CIvpeaster sp.; UF 54188; adoral view of test; Bermont Formation; lx. 


228 



fjrm 


Figure 3-36. Pleistocene irregular echinoids. 

A) Encope aberrans Martens, 1867; UF 42001; aboral view of test; Bermont 
Formation; 0.75x. 

B) Encope aberrans Martens, 1 867; UF 42001 ; adoral view of test; Bermont 
Formation; 0.75x. 

C) Encope michelini L. Agassiz, 1841; UF 62981, aboral view of test; Anastasia 
Formation; 0.5x. 

D) Encope michelini L. Agassiz, 1841; UF 62981, adoral view of test; Anastasia 
Formation; 0.5x. 


230 



Figure 3-37. Pleistocene irregular echinoids. 

A) Encope michelini L. Agassiz, 1841; UF 101080; aboral view of test; Bermont 
Formation; 0.75x 

B) Encope michelini L. Agassiz, 1841; UF 101080; adoral view of test; Bermont 
Formation; 0.75x 


232 




Figure 3-38. Pleistocene irregular echinoids. 

A) Mellita auinauiesperforata (Leske, 1778); UF 14778; aboral view of test; 
Satilla Formation; lx. 

B) Mellita auinauiesperforata (Leske. 1778); UF 14778; adoral view of test; 
Satilla Formation; lx. 


234 



Figure 3-39. Pleistocene irregular echinoids and Recent specimens for 
comparison. 

A) Moira atropos (Lamarck, 1 81 6); UF 1 001 80; aboral view of crushed test; 
Bermont Formation; lx. 

B) Moira atropos (Lamarck, 1816); UF 100180; adoral view of crushed test; 
Bermont Formation; lx. 

C) Moira atropos (Lamarck, 1816); UF 100180; lateral view of crushed test; 
Bermont Formation; lx. 

D) Moira atropos (Lamarck, 1816); UF 100180; posterior view of crushed test; 
Bermont Formation; lx. 

E) Moira atropos (Lamarck, 1816); UF 12641; aboral view of Recent test; lx. 

F) Moira atropos (Lamarck. 1816); UF 12641; adoral view of Recent test; lx. 

G) Moira atropos (Lamarck. 1816); UF 12641; lateral view of Recent test; lx. 

H) Moira atropos (Lamarck, 1816); UF 12641; posterior view of Recent test; lx. 


236 



237 


Class Crinoidea Fossils 
Lower Ocala Limestone Crinoids 

Class Crinoidea Miller, 1821 
Order Comatulida A.H. Clark, 1908 
Family Himerometridae A.H. Clark, 1908 
Genus Himerometra A.H. Clark, 1907 

Description: Centrodorsal low hemispherical to discoidal with concave to 
deeply depressed dorsal area. Cirrus sockets without distinct ornament, closely 
placed in two or three irregular marginal circles. Cirrals with or without dorsal 
spines. Ventral side of centrodorsal with interradial ridges and Y-shaped coelo- 
mic furrows. Basal rosette, but no rod-shaped basal rays in Recent species. 
Radials with a low free surface or concealed. Articular face steep. Interarticular 
ligament fossae very large, separated by wide midradial furrow. Ventral mus- 
cular fossae form narrow bands along ventral edge. Radial cavity large. Arms 
divided at primibrachs 2 and secundibrachs 4, exceptional at secundibrachs 2, 
and often at tertibrachs 2 of inner branches and tertibrachs 4 of outer branches. 
Proximal brachials narrow, laterally free and well separated. Pinnules from 
secundibrachs 2 and tertibrachs 2 larger than succeeding pinnules. Proximal 
pinnules may be carinate. The Eocene species H. bassleri Gislen differs in 
absence of coelomic furrows and presence of rod-shaped basals. In the 
Oligocene H. qrippae Anderson basals and coelomic furrows are unknown. 

Florida species: H. bassleri Gislen, 1934. 

Comments: The species is present in the Lower Ocala Limestone in 
peninsular Florida. Specimens consist of disarticulated ossicles including 
centrodorsals and brachials, as well as associated basal rays and radial plates. 


238 


Himerometra bassleri Gislen, 1934 
(Figure 3-40, A-C) 

Material examined: UF 39067 (figured centrodorsal), UF 39088 (figured 
centrodorsal), UF 39054-UF 39090 (50 centrodorsals, 53 radial plates, 20 basal 
rays). 

Formation; Lower Ocala Limestone. 

Locality; Inglis 01 A (CI001); Citrus County, FL; Yankeetown Quadrangle, 
SE1/4, SE1/4, Sec. 9, T17S, R16E. 

Collectors: FLMNFI crew. 

Date. 1974. 

Description: Centrodorsal a flattened hemisphere; ventral outline 
irregular pentagonal shaped. Dorsal surface partly cirrus free with a deep 
depression, indistinctly lobate radially. Cirri in two to three alternating whorls, 
about nine in each radial area. Cirrus sockets with a very indistinct tubercle on 
each side of the nerve lumen; no striation discernible. Proximal cirrals short, 
then increasing in length. Outer cirrals shorter, again becoming compressed 
laterally and with faint dorsal carination ending distally in slight eminence; 
eminence gradually develops into blunt tubercle when length of cirral has 
decreased to approximate square shape. Opposing spine well developed. 
Terminal claw long and curved. Dorsal surface of radials smooth, rather narrow, 
broader at interradial corners. Dorsal ligament pit twice as broad as the nerve 
lumen. Interarticular fossae much larger than muscle impressions, which are 
only narrow ventral bands; a deep radial notch is present between muscle 
fossae. Radial cavity medium-sized to large, sloping toward the central 


239 


depression with numerous septal ridges. No central calcareous plug. Basal 
impressions of centrodorsal rather distinct; radial portions of ventral side almost 
smooth; entire ventral surface slightly concave. Proportion between centrodorsal 
diameter and centrodorsal cavity approximately 0.21-0.27. Radials with dorsal 
facet toward centrodorsal are smooth; smallest radials and radial rings have 
fewer septal ridges on slopes facing central depression. Fixed primibrachial 
smooth, well rounded laterally. Secundaxillary with fixed primibrachial forming a 
very distinct synarthrial eminence. Pimaxil pentagonal in shape. Arm bases 
rather slender, well separated laterally. All division series of brachials smooth 
with indistinct synarthrial eminences. Number of arms varies from 35 to 45. 

Arms are smooth, rather slender, and proportionally long-jointed. Proximal 
pinnules rather stout and smooth. Basal segments are short and angular, slightly 
carinate, the distal a little longer. 

Upper Ocala Limestone Crinoids 

Fam., gen. et sp. incertae 
(Figure 3-40, D-G) 

Material examined: UF 48126 (figured centrodorsal), UF 48125 (2 
figured brachial plates). 

Formation: Upper Ocala Limestone. 

Locality: Wrights Creek (HO001); Holmes County, FL; Bonifay 
Quadrangle, SW1/4, SE1/4, Sec. 2, T5N, R15W. 

Collector: S. Burttschell. 


Date: 8/78. 


240 


Description: Centrodorsal pentagonal in horizontal outline; low dome or 
subhemispherical profile, broadly curving. Cirrus sockets in 3 horizontal rows, 
closely spaced with smallest sockets near dorsal apex. Ventral surface with 
moderate-sized radial cavity; opening approximately 0.4 of ventral surface 
diameter. Basal and radial plates, cirri, and arms not present on specimen. 

Comments: The fossils collected from Holmes County are identified as 
comatulid crinoids, but more precise taxonomic identification has not been 
attempted at this time. Very limited research is available regarding North 
American comatulids, and this also is true worldwide with respect to fossil 
comatulid crinoids. Therefore, documentation and reference materials (both as 
actual specimens and literature) are exceptionally difficult to obtain for 
comparison purposes. Review of the Treatise of Invertebrate Paleontology and 
the most thorough monograph series on modern crinoids written to date by A.H. 
Clark (i.e., Clark, 1915, 1921, 1931, 1941, 1947, 1950; Clark and Clark, 1967) 
resulted in no clear matches for these fossils. Therefore, I interpret the 
specimens to represent a new fossil species awaiting formal description. Herein 
the fossils are counted as a unique species of crinoid for the echinoderm 
diversity total. I consider them to be a new taxonomic record, but do not include 
this taxon as a new stratigraphic record since they have been reported earlier 
(Oyen, 1995). 


241 


Class Asteroidea Fossils 

Eocene Asteroids 

Class Asteroidea de Blainville, 1830 
Family Oreasteridae Fisher, 1911 
Genus Goniodiscaster H.L. Clark, 1909 

Description: Massive, flat body form with disc relatively large; 
pentagonal; arms typically 5, short; very broad at base, abactinal plates stellate, 
typically six-pointed, adradial facets of near radial ossicles highly elongate; some 
near radial ossicles transversely elongate. Actinals elliptical. 

Comments: One or more species may be present in the Ocala 
Limestone. Flowever, until more complete specimens are discovered no 
definitive taxonomic placement is possible. 

cf. Goniodiscaster sp. 

(Figures 3-40, H-J and 3-41, A-B) 

Material examined: UF 28135-UF 28137 (figured external molds of 
juveniles), UF 50000 (figured partial test), UF 38251 (figured marginal ossicle), 

UF 17244 (figured silicone peel of external mold and external mold of test). 

Formation: Ocala Limestone. 

Locality: Dickerson Limerock Mines (Flaile Complex) (AL004); Alachua 
County, FL, Newberry Quadrangle, T9S, R17E, Inglis 01A (CI001), Citrus 
County, Yankeetown Quadrangle, SE1/4, SE1/4, Sec. 9, T17S, R16E, Dolime 
Quarry 01 (CI009), Citrus County, Yankeetown Quadrangle, SE1/4, Sec. 11, 
T17S, R16E. 


242 


Collector; R. Portell and Jon Bryan (UF 28135-UF 28137), C. Oyen (UF 
50000), FLMNH Crew (UF 38251), R. Portell and Jon Bryan (UF 17244). 

Date. 03/18/88, 03/4/89, 1974, 3/18/88 (respectively). 

Description: UF 50000 - body form flat, disc large, length of major radius 
85 mm and length of minor radius 50 mm, arms 5, arms very broad at base; 
Superomarinals large and inflated, paired with long axes oriented perpendicular 
to arm margins; no visible pores; madreporite not visible; no spines present. UF 
17244 - abactinal ossicles stellate (six pointed). 

Comments; Specimens UF 28135-UF 28137 are external molds of 
juvenile asteroids which are more difficult to identify. However, their larger 
superomarginal ossicles help with taxonomic placement. A modern oreasterid 
species is shown for comparison purposes in Figure 3-42. Isolated ossicles are 
common in the Ocala Limestone and because of their large size, easy to find. 
Articulated specimens (whether molds or original material) are very rare. 
Specimens were found in the Upper and Lower Ocala Limestone. 

Oligocene Asteroids 

cf. Goniodiscaster sp. 

Material examined: UF 27464 (42 isolated ossicles). 

Formation: Suwannee Limestone. 

Locality: Terramar 01 (PO017), Polk County, Socrum Quadrangle, SI/4, 
Sec. 10, T26S, R22E. 


Collectors; R. Portell et al. 


243 


Date: 09/06/89 

Description: See Eocene section for description of cf. Goniodiscaster sp. 

Comments: Oligocene sea star ossicles are not nearly as common as 
those from the Ocala Limestone and they do not reach the large proportions as 
specimens of Eocene age. 

Miocene Asteroids 

Order Paxillosida Perrier, 1884 
Family Astropectinidae Gray, 1840 
Gen. et sp. incertae 

Material examined: UF 25333 (3 isolated superomarginal ossicles), UF 
32651 (superomarginal ossicle), UF 25132 (superomarginal ossicle). 

Formation: Parachucia Formation. 

Locality: White Springs (FIA001); Flamilton/Columbia counties. FL; W1/4, 
NW1/4, SW1/4, Sec. 7, T2S, R16E. 

Collectors: R. Portell and G. Morgan. 

Date: 12/13/89. 

Description: Marginal ossicles small; longer than wide (average length 4 
mm, average width 2 mm). 

Comments: Until better preserved material is found (possibly a partially 
articulated specimen) precise taxonomic assignment is problematic. A modern 
astropectinid species is shown for comparison purposes in Figure 3-42. 


244 

Pliocene Asteroids 

Order Forcipulatida Perrier, 1884 
Family Heliasteridae Viguier, 1878 
Genus Fleliaster Gray. 1840 

Description; Disc large, not set off externally from the fused bases of the 
rays, little elevated, with reticulated abactinal skeleton, and more or less 
numerous spines, pedicellariae, and papulae. Rays numerous, more than 20 in 
normal adults, more or less united at base, so that only a relatively small part 
(15-70%) is free. Adambulacral armature variable, usually single, sometimes 
double, especially near tip of ray; spines of alternate plates often of two sharply 
contrasted sizes, especially near base of ray. Pedicels arranged in two 
somewhat zigzag rows, so that near the middle of the ray they are distinctly 
quadriserial. Forcipate and forticate pedicellariae both present, the latter often of 
two distinct sizes. Interbrachial septa double and well developed, expanding at 
inner (proximal) end and uniting laterally, to form a discobrachial wall, so the 
cavity of the disc is almost completely separated from the ray cavities. 

Florida species: Fleliaster microbrachius Xantus, 1860. 

Comments: This species is present in the Tamiami Formation (Pliocene). 
Jones and Portell (1988) published the first fossil record of this species based 
upon these well-preserved fossils from southwestern Florida. The fossils are 
found within a matrix of quartz sand cemented with calcite and often are densely 
distributed as a result of imbrication of the sea stars. Jones and Portell (1988) 
reported finding the remains of over 360 individuals at the single locality in 
Charlotte County, with many individuals complete or nearly complete. 


245 


Heliaster microbrachius Xantus. 1860 
(Figure 43, A) 

Description: Rays total 27-44; rays more or less flattened abactinally, 
tapering rather sharply to a blunt point. Disc very large, some\A/hat elevated in 
well-preserved specimens, but not abruptly so. Abactinal skeleton stout, closely 
reticulated, with small meshes. Abactinal spines very numerous, 30-50 or more 
per square cm, small, usually low, more or less cylindrical and without definite 
arrangement. In some large specimens, spines show slight tendency to be 
capitate, and in many cases are very compressed. In some individuals spines on 
rays form five fairly distinct series, traceable inward for a variable distance onto 
the disc. At edge of disc the marginal series of adjoining rays sometimes very 
clearly separated by a bare space. Sides of ray with two series of compressed 
spines. Actinal surface very much as in H. helianthus , but pedicellariae are less 
frequent and reduction of the adambulacral armature reaches its extreme; in 
large specimens only every other adambulacral plate bears a spine until the 
distal half or third of furrow is reached, and even at extreme tip of the ray it is rare 
to find a plate with two spines. Pedicels numerous, distinctly quadriserial at 
middle of ray. Madreporite rather small, often concave, and usually fragmented. 


Order Paxillosida Perrier, 1884 
Family Astropectinidae Gray, 1840 
Gen. et sp. incertae 

Material examined: UF 48093 (2 superomarginal ossicles). 


Formation: Tamiami Formation. 


246 


Locality: Lomax-King Pit A (CH028), Charlotte County, FL; Punta Gorda 
Quadrangle, SW1/4, SE1/4, Sec.28, T41S, R23E. 

Collector: R. Portell. 

Date: 12/23/88 

Description: Marginal ossicles small; longer than wide. Intermarginal 
facet well-defined. 

Comments: Until better preserved material is found (possibly a partially 
articulated specimen) precise taxonomic assignment is problematic. 

Order Paxillosida Perrier, 1884 
Family Luidiidae Sladen, 1899 
Genus Luidia Forbes, 1839 

Description: Arms long, flat, strap-like, numbering from a minimum of 
five total to several more (undefined). Small central disc; absence of 
superomarginal plates; paxillar abactinal surface and elongate inferomarginal 
plates extending from the ambitus to the adambulacral plates. Inferomarginal 
ossicles with a conspicuous fringe of inferomarginal spines. 

Florida species: Luidia sp. 

Comments: This genus is represented by an incomplete specimen of a 
species waiting formal description. The fossil is from the Tamiami Formation and 
was collected in association with the Fleliaster microbrachius described herein. 

Luidia sp. 

(Figure 3-43, B) 

Material examined: UF 60184 (one incomplete test). 


247 


Formation; Tamiami Formation. 

Locality; El Jobean 01 (CH002); Charlotte County, FL; El Jobean 
Quadrangle, SW1/4, NE1/4, Sec. 21, T40S, R21E. 

Collector; D. Swanson. 

Date; Unknown. 

Description; Five-armed asteroid; aboral surface with two partial arms; 
portion of central disk visible but distorted. Major radius at least 46.5 mm, minor 
radius 14.7 mm. Arms elliptical, strap-like, disk rather flat, small; arm taper 
gradual; madreporite nor spines visible. 

Comment; This is the first report of the genus from the Cenozoic of the 
southeastern United States. A nearly complete test of this taxon is currently in 
the hands of a private collector. Hopefully it will be donated to a museum for 
formal description. 


Class Ophiuroidea Fossils 

Eocene Ophiuroids 

Class Ophiuroidea Gray, 1840 
Order Ophiurida Muller and Troschel, 1840 

Florida species; All identifications limited to taxonomic order at this time, 

with only one tentative proposal at the familial level. 

Comments; Most fossils collected thus far consist of disarticulated 

vertebral ossicles, thereby limiting the potential for lower level identifications. 

Ophiurioid taxonomy is based mainly on the features of the disk and the gross 

anatomy of complete specimens. Currently, no reference exists for identification 


248 


of disarticulated vertebral ossicles. Ophiuroids are present in the Avon Park 
Formation (Eocene), Marks Head Formation (Miocene), and Tamiami Formation 
(Pliocene) in Florida. 


cf. Amphiuridae 
Gen. et sp. indet. 

(Figure 3-40, H) 

Material examined; UF28129-28130 (figured external molds of juveniles 
on Thalassodendron seagrass blades. 

Formation: Avon Park Formation. 

Locality: Dolime Quarry 01 A (CI013); Citrus County, FL; Yankeetown 
Quadrangle, SE1/4, Sec. 11, T17S, R16E. 

Collectors R. Portell and J. Bryan. 

Date: 03/18/88. 

Description; Individuals with five arms that are long and thin with short 
erect spines. 

Comments: UF 28129-28130 are external molds of juvenile specimens 
which are more difficult to identify. Specimens have been found only associated 
with sea grass fossils in the Avon Park Formation. 

Miocene Ophiuroids 

Gen. et sp. indet. 

(Figure 3-43, C) 

Material examined: UF 45853 (figured vertebral ossicle). 

Formation: Marks Head Formation. 


249 


Locality; Brooks Sink (BF001); Bradford County, FL; Brooker 
Quadrangle, SW1/4, SW1/4, Sec. 12, T7S, R20E. 

Collector: R. Portell. 

Date: 01/14/88. 

Description; Four disarticulated vertebral ossicles; small (one to two mm 
diameter); micromorphological structures such as vertical dumbbell, dorsal 
muscle attachment, ventral muscle attachment, ventral nose, and podial basin 
are preserved and visible. 

Comments: The fossil ophiuroid ossicles from Brooks Sink are well- 
preserved, but due to the limited number of skeletal components, identification to 
lower taxonomic levels is not possible. The fossils are important, however, 
because they prove that ophiuroids have a fossil record in Florida, and with 
detailed examination of the fine-fraction of disaggregated limestones it is possible 
to recover such taxa. For the purpose of biostratigraphic analysis in this 
dissertation, these fossils represent a new stratigraphic record for the Marks 
Flead Formation, but cannot be considered as a new taxonomic record without 
additional, more complete material for identification. 

Pliocene Ophiuroids 

Gen. et sp. incertae sedis 
(Figure 3-43, D) 

Material examined: UF 7380 (complete but recrystallized test). 


Formation: Tamiami Formation. 


250 


Locality: El Jobean 01 (CH002); Charlotte County, FL; El Jobean 
Quadrangle, SW1/4, NE1/4, Sec. 21, T40S, R21E. 

Collector: R. Portell. 

Date: 4/86. 

Description: Arms five, generally subcircular in outline, narrowing toward 
the distal termination; short conical spines on arms; disc covered by fine 
imbricating scales and minute spines; specimen recrystallized preventing more 
detailed description. 

Comments: UF 7380 is preserved on the sun star Fleliaster 
microbrachius . It is very small (up to three cm or slightly less in diameter with 
arms extended). The detailed morphology of this ophiuroid is obscured by 
recrystallization of calcite and by surficial cemented matrix. The combination of 
small size and attached matrix prevents more precise taxonomic identification of 
the fossils. Flerein, I do not consider the identification to reflect a new 
stratigraphic record or a new taxonomic record based solely on these fossils, but 
rather only note these as an occurrence of the class Ophiuroidea from the 


Tamiami Formation. 


Figure 3-40. Eocene crinoids, asteroids, and ophiuroids. 

A) Himerometra bassleri Gislen; UF 39067; dorsal view of centrodorsal element; 
Ocala Limestone; 3x. 

B) Flimerometra bassleri Gislen; UF 39067; ventral view of centrodorsal element 
with attached radial plates; Ocala Limestone; 3x. 

C) Flimerometra bassleri Gislen; UF 39088;ventral view of centrodorsal element 
with radial plates removed (allowing distinctive rod-shaped basal rays to be 
observed); Ocala Limestone; 3x. 

D) Unidentified comatulid crinoid; UF 48126; dorsal view of centrodorsal 
element; Ocala Limestone; 7x. 

E) Unidentified comatulid crinoid; UF 48126; ventral view of centrodorsal 
element; Ocala Limestone; 7x. 

F) Unidentified comatulid crinoid; UF 48125; brachial plate; Ocala Limestone; 7x. 

G) Unidentified comatulid crinoid; UF 48125; brachial plate; Ocala Limestone; 

7x. 

FI) cf. Goniodiscaster sp. and cf. Amphiuridae gen. et sp. indet.; UF 28135- 
28137 (asteroids) and UF281 29-281 30 (ophiuroids); external molds of 
unidentified juveniles among Thalassodendron auricula-leporis seagrass 
blades; Avon Park Formation; 2x. 

I) cf. Goniodiscaster sp.; UF 50000; view of specimen showing marginals and 
abactinal ossicles; Ocala Limestone; lx. 

J) cf. Goniodiscaster sp.; UF 38251 ; lateral view of marginal ossicle; Ocala 
Limestone; 2x. 


252 





Figure 3-41. Eocene asteroid and silicone rubber peel. 

A) cf. Goniodiscaster sp.; UF 17244; RTV silicone rubber peel of incomplete 
test; Ocala Limestone; lx. 

B) cf. Goniodiscaster sp.; UF 17244; external mold of incomplete test; Ocala 
Limestone; lx. 




254 





Figure 3-42. Recent asteroids illustrating articulated ossicle arrangements that 
may be similar to taxa in Florida’s fossil record. 

A) Oreaster reticulatus (Linnaeus, 1758); UF1 05550; adoral view of Recent test, 
collected offshore near Tampa, FL, showing large marginal ossicles; lx. 

B) Astropecten sp.; UF30969; aboral view of Recent test, collected from the Gulf 
of Mexico, showing marginal ossicles; 1x. 

C) Astropecten sp.; UF30969; adoral view of Recent test, collected from the Gulf 
of Mexico, showing marginal ossicles; lx. 


256 




Figure 3-43. Pliocene asteroids along with Miocene and Pliocene ophiuroids. 

A) Heliaster microbrachius Xantus; UF 7423; top view of block with over 12 
individuals preserved in sandstone; Pliocene, Tamiami Formation, 0.3x. 

B) Luidia sp.; UF 60184; top view of two nearly complete rays, adhered to 
Fleliaster microbrachius block; Pliocene, Tamiami Formation; lx. 

C) Ophiuroid; UF lot 45853; vertebral ossicle; Miocene, Marks Ftead Formation; 
7x. 

D) Unidentified ophiuroid; UF 7380; specimen nestled among rays of Heliast ^ 
microbrachius ; Pliocene, Tamiami Formation; 1.5x. 


258 



CHAPTER 4 

ECHINODERM DIVERSITY PATTERNS AND BIASES 

Taxonomic and Biostratigraphic Discussion 

Several points should be noted about the data presented in this paper \A/ith 
respect to taxonomic and stratigraphic records. Complete tests, test fragments, 
spines, and molds were included in the echinoid database, as well as articulated 
and disarticulated ossicles, and some molds of crinoids, asteroids, and 
ophiuroids are part of my records. New stratigraphic records include any 
echinoderms reported herein that have not been previously published elsewhere 
when referring to that particular stratigraphic unit. This may have some bias 
associated with it due to stratigraphic nomenclature changes, and it is discussed 
in more detail in a later section on biases in the Florida echinoderm record. The 
taxonomic records include old and new (but not yet formally described) taxa. I 
have not counted subspecies as unique taxa because, in my opinion, most of 
them need to be re-examined for possible synonymy. Additional specific notes 
regarding newly reported taxa are included in the systematic paleontology 
chapter earlier in this dissertation. Finally, most of the uncertain records (both 
taxonomic and stratigraphic) were omitted to provide a conservative and reliable 
database for presentation in this paper. 


259 


260 


Eocene Echinoids 

Fossil echinoids from the Eocene in Florida are reasonably well-known 
and diverse. A variety of fossils from the carbonates of the Middle and Upper 
Eocene have been collected by paleontologists for more than 150 years. 
Consequently, most of the Eocene taxa and biostratigraphic records are not new. 
When compared with other epochs, the Eocene shows little change 
(proportionally) in terms of new stratigraphic occurrences or potential new taxa. I 
interpret this to be a result of the many years of intensive collecting. A second 
factor, which may contribute to the relatively small number of new Eocene 
echinoid records, is the good preservation of the calcitic tests in the carbonates. 
These tend to be preserved as complete or nearly complete specimens, unlike 
many echinoids of the siliciclastic-rich Neogene units. The preservation bias will 
be discussed in more detail later. 

The Eocene exhibits the highest number of taxa of any epoch from the 
Middle Eocene through the Pleistocene. The proportion of echinoid taxa during 
this epoch is three (= 7.9%) regular echinoids to 35 (= 92.1%) irregular echinoids, 
bringing the total number to 38 (Figure 4-1). 

The regular echinoids are represented by the genera Phvllacanthus and 
Dixieus , as well as fragments and spines of another taxon that is a diadematoid. 
Of the regular echinoids, Phvllacanthus is most abundant, yet specimens are 
relatively rare as complete or nearly complete tests. All the regular echinoid taxa 
are found in the Upper Ocala Limestone while only Phvllacanthus is also present 
in the Lower Ocala Limestone. In many cases, the regular echinoid fossils are 


261 


FLORIDA ECHINOID DIVERSITY 



Figure 4-1 . Irregular and regular echinoid diversity reflecting updated values as a 
result of this study. Data are provided at epoch level resolution, and 
the light stipple pattern corresponds to irregular echinoid species 
while the black pattern represents regular echinoid species. 


broken or disarticulated, and may be represented by only plate fragments, 
radioles, or Aristotle’s lantern components. 

The Eocene irregular echinoids are the most diverse segment of the fossil 
echinoderm record of Florida. Twenty-two genera (Table 4-1) are found in rocks 


262 


from the Avon Park, Lower and Upper Ocala Limestone (see Figure 2-1, chapter 
2). Several genera, including Oliqopyqus . Rhvncholampas . Schizaster , and 
Eupataqus . have three to four species each and thereby are more species-rich 
than most of the other Eocene genera. The irregular echinoids generally have 
better preservation than the regular echinoids, though this is typical of 
preservation comparisons between the two groups throughout their entire fossil 
record and is not unique to the Eocene or to Florida echinoids (Kier, 1977; 
Donovan, 1991; Greenstein, 1993). Although there are numerous taxa, less than 
half of the species are common, and are easily found while conducting fieldwork. 
The four genera listed above, along with Weisbordella , Durhamella . 

Neolaqanum . and Periarchus (see Figures 3-1 through 3-4, chapter 3), are 
common constituents of the Florida Eocene. 

One group of echinoids has been used as an important stratigraphic tool 
for the Late Eocene. Zachos and Shaak (1978) developed a biostratigraphic 
scheme based on distribution of the three species of Oliqopyqus echinoids found 
in Florida, O. phelani Kier, O. haldemani (Conrad), and O. wetherbvi de Loriol 
(Figure 3-1). These echinoids work as good biostratigraphic markers because 
they are well-preserved, abundant, and easy to identify in the field. The species 
O. phelani is restricted to the Lower Ocala Limestone, with both O. haldemani 
and O. wetherbvi confined to the Upper Ocala Limestone. Most facies of the 
Ocala Limestone contain these echinoids and they are useful index fossils for 


field studies of the Middle to Late Eocene limestones of Florida. 


263 


Table 4-1 . Eocene echinoderms and the formations from which the fossils were 
collected. 


GENUS 

# OF TAXA 

FORMATION(S) 

ECHINOIDS 

Phyllacanthus 

1 

Upper Ocala Ls 
Lower Ocala Ls 

Dixieus 

1 

Upper Ocala Ls 

Diadematoida 

1 

Upper Ocala Ls 

Oligopygus 

3 

Upper Ocala Ls 
Lower Ocala Ls 

Amblypygus 

1 

Upper Ocala Ls 

Fibularia 

1 

Upper Ocala Ls 
Lower Ocala Ls 

Periarchus 

1 

Lower Ocala Ls 

Protoscutella 

1 

uncertain 

Weisbordella 

2 

Upper Ocala Ls 

Durhamella 

2 

Upper Ocala Ls 

Neolaganum 

2 

Upper Ocala Ls 
Avon Park Fm 

Wythella 

1 

Upper Ocala Ls 

Eurhodia 

1 

Upper Ocala Ls 

Echinolampas 

1 

Upper Ocala Ls 

Rhyncholampas 

4 

Upper Ocala Ls 
Lower Ocala Ls 

Schizaster 

3 

Upper Ocala Ls 

Ditremaster 

1 

Upper Ocala Ls 

Agassizia 

2 

Upper Ocala Ls 
Lower Ocala Ls 

Macropneustes 

1 

Upper Ocala Ls 

Brissopsis 

2 

Upper Ocala Ls 

Plagiobrissus 

2 

Upper Ocala Ls 
Lower Ocala Ls 

Eupatagus 

3 

Upper Ocala Ls 
Lower Ocala Ls 

Mortonella 

1 

Upper Ocala Ls 

CRINOIDS 

Himerometra 

1 

Lower Ocala Ls 

gen. indet. 

1 

Upper Ocala Ls 

ASTEROIDS 

Oreasteridae or 
Goniasteridae 

7 

Upper Ocala Ls 
Lower Ocala Ls 
Avon Park Fm 

OPHIUROIDS 

gen. indet. 

? 

Avon Park Fm 


264 


The stratigraphic distribution of echinoids in the Middle Eocene Avon Park 
Formation is limited to one species of irregular echinoid, Neolaqanum dalli . 
Twitchell (Figure 3-3), and no new stratigraphic records or new taxonomic 
records are reported for this formation. The Lower Ocala Limestone contains 10 
species (one regular and nine irregular echinoid species), with no new taxonomic 
or stratigraphic records from the unit. All of the changes in the Eocene data are 
found in the Upper Ocala Limestone. Within this unit, 29 species are present 
(three regular and 26 irregular echinoid species), and I report five new 
stratigraphic records and three new taxonomic records for this formation. 

An interesting aspect of the echinoid stratigraphic distributions in Florida is 
the relatively limited range of species. This likely is an artifact of the stratigraphic 
nomenclature itself within the state, since many of the formations were named (at 
least in part) on the basis of fossils. This will be addressed in more detail in 
following sections of this chapter, but at this point I will note which taxa are 
present in more than one stratigraphic unit (in the Eocene I consider the Upper 
Ocala Limestone and the Lower Ocala Limestone to be unique). This 
characteristic is relatively unusual for echinoids in Florida, with only three taxa 
that have this stratigraphic distinction in the Eocene. These are: Phvllacanthus 
mortoni (Conrad) (Lower and Upper Ocala Limestone), Fibularia vauqhani 
(Twitchell) (Lower and Upper Ocala Limestone), and Plaqiobrissus curvus 
(Cooke) (Lower and Upper Ocala Limestone). 


265 


Oligocene Echinoids 

The Oligocene is at the lower end of the spectrum of echinoid species 
diversity in the Florida fossil record. A dramatic decrease in the number of taxa 
from the Eocene to the Oligocene is obvious in the pattern of species diversity at 
that time (Figure 4-2). The Oligocene carbonates contain 1 1 total taxa, with the 
proportion of regular to irregular echinoids at three (= 27.3%) to eight (= 72.7%) 
respectively (Figure 4-1). Therefore, this total diversity represents a considerable 
drop from 38 taxa in the Eocene. The origin of this dramatic decrease in diversity 
is not well defined yet, but a similar pattern has been noted globally for both 
terrestrial and marine organisms (see Prothero and Berggren, 1992). One 
attempt to identify the cause of the echinoid diversity decrease from the Eocene 
to the Oligocene was by McKinney and Oyen (1989). They proposed that a 
temperature decrease more strongly correlated with the diversity drop than a 
lowering of sea level, which also matched the cause proposed for the Gulf Coast 
mollusk diversity drop (Dockery, 1986). Later, McKinney et al. (1992) expanded 
the focus and examined the global echinoid pattern across the Eocene- 
Oligocene boundary. They found a similar pattern in the global echinoid data as 
existed in the Gulf Coast region, and again proposed temperature decrease to be 
the cause of the diversity drop. This pattern is discussed in more detail in 
subsequent paragraphs of the chapter. 


266 


FLORIDA ECHINOID DIVERSITY 



Figure 4-2. Echinoid diversity curves for Florida. Open diamonds represent 
previously published taxonomic records while the black squares 
show the updated diversity values. Note distinct change in diversity 
trend from the Oligocene to the Miocene as a result of the data 
presented herein. 


The regular echinoid species of the Oligocene are members of two 
genera, Gaqaria and Phvmotaxis . Though none of the species are common, 
Gaqaria mossomi (Cooke) tends to be more prevalent than the other regular 
echinoids (Figure 3-9). All of the Oligocene regular echinoids known from Florida 
are found in the Suwannee Limestone. The typical pattern of fragmented and 
poorly preserved regular echinoids holds true for Oligocene taxa. 


267 


The irregular echinoids are represented by species from five genera 
(Table 4-2). The most diverse genus is Clypeaster , which has four species, and 
fossils from this genus are found in all three of the echinoid-bearing Oligocene 
formations in Florida. The most common species is C. roqersi (Morton), which 
occurs in both the Marianna Limestone of northern Florida and the Suwannee 
Limestone (Figure 3-13). The most pervasive and abundant echinoid in the 
Oligocene is Rhvncholampas qouldii (Bouve), which is present in the Suwannee 
Limestone throughout most of the unit’s areal extent (Figure 3-13). This species 
normally is well-preserved, easily identified in the field, and occurs in lithofacies 
varying from wackestones to grainstones (though more common in the sandier, 
higher energy end of the facies spectrum). 

Within the Oligocene, the 11 taxa I report herein are distributed among 
three formations (Figure 2-1). The Suwannee Limestone is the most diverse 
stratigraphic unit, with all 1 1 taxa found in the formation. I present five new 
stratigraphic records for the formation and two new taxonomic records. The 
Marianna Limestone has two species of echinoids, with one new stratigraphic 
record and no new taxonomic records. The Bridgeboro Limestone also has a 
total of two taxa, neither represents a new stratigraphic record nor a new 
taxonomic record for the state. 

Finally, four taxa are found in more than one formation of Oligocene age, 
which is proportionally high for Florida echinoids. These include: Clypeaster 
cotteaui Egozcue (Suwannee Limestone and Bridgeboro Formation), C. roqersi 
(Morton) (Suwannee Limestone and Marianna Limestone), Aqassizia mossomi 


Table 4-2. Oligocene echinoderms and the formations from which the fossils 
were collected. 


268 


GENUS 

# OF TAXA 

FORMATION(S) 

ECHINOIDS 

Gagaria 

1 

Suwannee Ls 

Phymotaxis 

2 

Suwannee Ls 

Clypeaster 

4 

Bridgeboro Ls 

Rhyncholampas 

1 

Marianna Ls 
Suwannee Ls 
Suwannee Ls 

Agassizia 

1 

Bridgeboro Ls 

Schizaster 

1 

Suwannee Ls 
Marianna Ls 

Brissopsis 

1 

Suwannee Ls 
Suwannee Ls 

CRINOIDS 

none 


ASTEROIDS 

Oreasteridae 

7 

Bridgeboro Ls 

OPHIUROIDS 

none 

Suwannee Ls 


Cooke (Suwannee Limestone and Bridgeboro Formation), and Schizaster 
americanus Clark (Suwannee Limestone and Marianna Limestone). 

Miocene Echinoids 

The Miocene shows the greatest change from previously published 
information on echinoid diversity (Figure 4-2). The increase in new taxa (11) and 
new stratigraphic records (16) is dramatic and represents a modification in the 


269 


overall pattern seen throughout the Cenozoic strata of Florida. Prior to my 
revisions in taxonomic and stratigraphic records, the diversity trend for echinoids 
in the state (as well as throughout the Gulf Coastal Plain) showed a marked drop 
from the Oligocene into the Miocene. An example of this pattern can be seen at 
a slightly broader scale for the southeastern U.S. and Caribbean echinoids in 
work done by McKinney and Oyen (1989) and McKinney et al. (1992). However, 
this drop in diversity was anomalous when compared with the global pattern, 
which shows an increase in diversity from the Oligocene to the Miocene (Kier, 
1977; McKinney et al., 1992). I believe this large increase in the known 
distribution of Florida echinoids is strongly a function of collector bias, because 
many of the additions I have made are based on incomplete specimens or poorly 
preserved (moldic) specimens, which many collectors tend to disregard (biases 
discussed below). The diversity data, therefore, fit the global echinoid pattern 
much more closely following my revision. The siliciclastic and carbonate units of 
the Miocene contain a total of 19 taxa, with 7 of these regular echinoids (= 

36.8%) and 12 irregular echinoids (63.2%). 

The echinoids of the Miocene represent the highest proportion of regular 
to irregular echinoid taxa from the Eocene through the Pleistocene. Four genera 
of regular echinoids are present in the Miocene, including Gaqaria , 
Psammechinus . Arbia . and Prionocidaris . Most of the regular echinoids are 
uncommon, but with closer examination of sediment samples such fragments are 
being recovered with increasing frequency. One exception to this pattern is 
Prionocidaris . whose test plates and radioles are common at most Chipola 


270 


Formation localities. Two examples of this incomplete (moldic) preservation can 
be seen in the photos of Figure 3-14 (in chapter 3). 

A total of at least eight genera of irregular echinoids are present in 
Florida’s Miocene units (Table 4-3). Most of these echinoids are not common. 
The best examples of locally concentrated specimens include internal and 
external molds of Lovenia dark! (Lambert) (Chattahoochee Formation) in 
Jackson County, and internal molds and tests of Abertella aberti (Conrad) 
(Coosawhatchie, Arcadia, and Peace River formations) from several locations in 
the state (Figures 3-16 and 3-18). Preservation of the Miocene echinoids rarely 
is complete and therefore identification to lower taxonomic levels is more difficult. 
Thus, my close examination of incomplete specimens partially accounts for the 
diversity increase in the Miocene. Since my taxonomic record data have been 
culled to include only specimens that are likely to be described as new species, 
and several fossils were omitted pending more detailed preparation and 
examination, I believe the number of taxonomic records will increase even more 
as my work continues. 

The echinoids from this epoch are found in nine of the ten echinoderm- 
bearing formations (Figure 2-1). The Chipola Formation has the greatest number 
of taxa (eight total), including several which may be undescribed species. 
Recently, Rhvncholampas chipolanus Oyen and Portell was described from the 
Chipola Formation based on a single, well-preserved specimen (Figure 3-16). 
Since that description, additional test fragments from other individuals of this 


271 


Table 4-3. Miocene echinoderms and the formations from which the fossils were 
collected. 


GENUS 

# OF TAXA 

FORMATION(S) 

ECHINOIDS 

Gagaria 

2 

Parachucia Fm 

Arbia 

1 

Chattahoochee Fm 

Psammechinus 

1 

Chipola Fm 

Prionocidaris 

1 

Chipola Fm 
Torreya Fm 

gen. indet. 

3 

Arcadia Fm 
Chattahoochee Fm 
Shoal River Fm 

Mellitidae 

1 

Statenville Fm 

Abertella 

2 

Arcadia Fm 
Chipola Fm 
Coosawhatchie Fm 
Peace River Fm 
Torreya Fm 

Clypeaster 

2 

Chattahoochee Fm 
Chipola Fm 

Rhyncholampas 

3 

Arcadia Fm 
Chipola Fm 
Peace River Fm 

Echinocyamus 

1 

Chipola Fm 

Echinarachnius 

1 

Chipola Fm 

Agassizia 

1 

Arcadia Fm 

Brissopatagus 

1 

Chattahoochee Fm 

Brissidae 

1 

Chipola Fm 

Lovenia 

1 

Chattahoochee Fm 

gen. indet. 

1 

Coosawhatchie Fm 

CRINOIDS 

none 


ASTEROIDS 

Astropectinidae 

? 

Arcadia Fm 
Chipola Fm 
Coosawhatchie Fm 
Parachucia Fm 

OPHIUROIDS 

gen. indet. 

? 

Marks Head Fm 


272 


species have been located in samples from the Chipola Formation donated to the 
FLMNH by Emily and Harold Vokes. Two other formations (the Arcadia and 
Chattahoochee) have four and six different taxa (respectively), while the rest of 
the formations only have one or two species. Within the Miocene, a total of 19 
taxa are present, with 11 of those as new taxonomic records. In addition, 16 new 
stratigraphic records for the state are presented herein. The Chattahoochee 
Formation shows the most change of the nine stratigraphic units containing 
echinoids, with six new stratigraphic records and five new taxa. 

The Miocene has four taxa that are found in more than one stratigraphic 
unit. These include: Prionocidaris cookei Cutress (Chipola and Torreya 
formations), Abertella aberti (Arcadia, Coosawhatchie, and Peace River 
formations), Abertella sp. (Chipola, Coosawhatchie, and Torreya formations), and 
Rhvncholampas sp. (Arcadia and Peace River formations). 

Pliocene Echinoids 

The Pliocene echinoid record shows modest change with respect to the 
number of taxa hitherto known from Florida, but has a significant number of new 
stratigraphic records. Perhaps one reason for this is the significant urban 
development in the areas underlain by Pliocene sediments, especially in areas of 
south Florida. As excavation of land takes place, new (and typically temporary) 
pits are produced which expose fossiliferous strata for examination. This factor, 
along with stratigraphic nomenclature revisions, may have enhanced the 


273 

potential for additional taxa. Currently, I have identified 23 taxa from the 
Pliocene, with five regular (= 21.7%) and 18 irregular echinoids (= 78.3%). 

Four genera of regulars are found in the Pliocene, including Lytechinus , 
Echinometra , and Eucidaris represented by a single species, and Arbacia 
potentially having up to four species (see Table 4-4). Preservation is relatively 
good in specimens of Lytechinus . Echinometra , and Arbacia . Specimens of the 
species Lytechinus varieqatus (Lamarck) and Echinometra lucunter (Linnaeus) 
are very well-preserved, showing little diagenetic or compaction effects (Figure 3- 
19). Flowever, all specimens of Pliocene Eucidaris tribuloides (Lamarck) 
collected thus far show extensive compaction and fragmentation (Figure 3-19). 

At least nine genera (represented by up to 18 species) of irregular 
echinoids also are present. Most Pliocene genera are represented by only one 
or two species, but the clypeasteroids exhibit slightly greater diversity. The 
genus Clvoeaster contains at least five species from four formations and Encope 
has two species from four formations. The other genera of irregular echinoids 
with more than one species include Rhvncholampas . Mellita . and Pericosmus , 
with each having two. Preservation of the irregular echinoids generally is good to 
excellent, though some beds have only fragmentary remains. Several species of 
these irregular echinoids can be found in high concentration locally. Examples of 
common and easily recognizable Pliocene echinoids from Florida include the 
mellitids Encope tamiamiensis Mansfield (Figures 3-27 and 3-29) and Mellita 
aclinensis Kier from the Tamiami Formation. These two species were found in 


such high concentrations in some locations that the tests were densely 


274 

imbricated (e.g., UF locality CH003, the former Lomax King Pit in Charlotte 
County). 

The echinoids from the Pliocene are present in five formations (Figure 2-1) 
and have 23 taxa of regular and irregular echinoids (Table 4-4). The Tamiami 
Formation has the highest diversity of the stratigraphic units, \A/ith 12 taxa 
present. The Caloosahatchee Formation (seven species) and the Nashua 
Formation (up to six taxa) follow the Tamiami Formation in diversity, with the 
Intracoastal Formation containing three taxa. A large number of new 
stratigraphic records (15) of echinoids are reported herein and the unit with the 
largest number of new records is the Nashua Formation (five records). The 
Tamiami Formation also has a significant number of additions, with three new 
stratigraphic records. All formations in the Pliocene, except the Caloosahatchee, 
have new records of echinoids. Three formations have possible new species , 
but each are quite tentatively considered new at this time. 

In the Pliocene, five species are found in more than one stratigraphic unit. 
These include: Lytechinus varieqatus (Lamarck) (Caloosahatchee and Tamiami 
formations), Arbacia improcera (Conrad) (Tamiami and Jackson Bluff 
formations), Encope aberrans Martens (Caloosahatchee, Tamiami, and 
Intracoastal formations), Aqassizia porifera (Ravenel) (Caloosahatchee and 
Tamiami formations), and Echinocardium orthonotum (Conrad) (Tamiami, 
Intracoastal, and Jackson Bluff formations). 


275 


Table 4-4. Pliocene echinoderms and formations from which they were collected. 


GENUS 

# OF TAXA 

FORMATION(S) 

ECHINOIDS 

Lytechinus 

1 

Caloosahatchee Fm 
Tamiami Fm 

Echinometra 

1 

Caloosahatchee Fm 

Arbacia 

2 

Nashua Fm 
Jackson Bluff Fm 
Tamiami Fm 

Eucidaris 

1 

Tamiami Fm 

Encope 

2 

Caloosahatchee Fm 
Intracoastal Fm 
Nashua Fm 
Jackson Bluff Fm 
Tamiami Fm 

Mellita 

2 

Nashua Fm 
Tamiami Fm 

Clypeaster 

5 

Caloosahatchee Fm 
Intracoastal Fm 
Nashua Fm 
Tamiami Fm 

Rhyncholampas 

2 

Caloosahatchee Fm 
Tamiami Fm 

Pericosmus 

2 

uncertain 

Agassizia 

1 

Caloosahatchee Fm 
Tamiami Fm 

Echinocardium 

1 

Intracoastal Fm 
Nashua Fm 
Jackson Bluff Fm 
Tamiami Fm 

Plagiobrissus 

1 

Tamiami Fm 

Brissidae 

1 

uncertain 

CRINOIDS 

none 


ASTEROIDS 

Heliaster 

1 

Tamiami Fm 

Luidia 

1 

Tamiami Fm 

Oreasteridae 

7 

Tamiami Fm 

gen. indet. 

? 

Nashua Fm 

OPHIUROIDS 


? 

Tamiami Fm 


276 


Pleistocene Echinoids 

The Florida Pleistocene echinoid record has doubled since I began my 
work to update the echinoderm biostratigraphy in the early 1990’s. The diversity 
now stands at six taxa, which is interesting not only due to the large proportional 
increase, but also because of the very low diversity overall during the epoch. 

The drop from the Pliocene to the Pleistocene is dramatic, even with my 
increased diversity values (Figure 4-2). Furthermore, the Pleistocene diversity is 
much lower than the 27 species known from the shallow water environments 
(<37 m) surrounding Florida today (Camp et al., 1998). The strata of the 
Pleistocene contain no regular and six irregular echinoids. This is the only epoch 
in Florida currently without any regular echinoids (Figure 4-1). 

The irregular echinoids are represented by four genera (Table 4-5), with 
Encope and CIvpeaster each having two species while Mellita and Moira have 
one species each. Preservation of echinoids in the Late Pleistocene units is 
relatively poor, but in Early Pleistocene formations the preservation is commonly 
better. This good preservation is obvious in species like CIvpeaster rosaceus 
(Linnaeus), shown in Figure 3-35 of chapter 3. Many Pleistocene samples are 
fragmented and heavily abraded, which may reflect a high energy, near-shore 
depositional environment. In some cases, the samples have sedimentary rock 
and other fossils well-cemented to their surfaces, as in Encope michelini Agassiz 
(Figure 3-36), making them difficult to clean, prepare, and identify. 

The Pleistocene echinoids are part of three formations in the state (Figure 
2-1). The Bermont Formation has four taxa, the Anastasia Formation 


277 


Table 4-5. Pleistocene echinoderms and the formations from which the fossils 
were collected. 


GENUS 

# OF TAXA 

FORMATION(S) 

ECHINOIDS 

Encope 

2 

Anastasia Fm 
Bermont Fm 

Mellita 

1 

Anastasia Fm 
Satilla Fm 

Clypeaster 

2 

Bermont Fm 

Moira 

1 

Bermont Fm 

CRINOIDS 

none 


ASTEROIDS 

gen. indet. 

9 

Bermont Fm 

OPHIUROIDS 

none 



two taxa, and the Satilla Formation one taxon. Four new stratigraphic records 
are found in the Pleistocene, with the Bermont Formation having two, and each 
of the other formations having one. Only one new taxonomic record resulted 
from my work, which was found in the Bermont Formation. 

Finally, only one species is found in more than one formation during this 
epoch. The taxon is Mellita quinquiesperforata . and it is found in the Anastasia 
and Satilla formations. 

Crinoids 

The first report of Eocene comatulid crinoids in North America was by 
Emmons (1858), in which he described a new species, Microcrinus conoideus . 


278 


from the Eocene marls of North Carolina. A second crinoid worker, Gislen 
(1934), described the comatulid Himerometra bassleri from the Eocene of South 
Carolina. Howe (1942) published a discussion of Tertiary microfossils that he 
believed had been overlooked by Gulf Coast paleontologists, even though the 
fossils may be locally abundant. At the time, Howe was unable to find prior 
references to four classes of fossil echinoderms, including ophiuroids, comatulid 
crinoids, asteroids, and holothurians, from strata in the Gulf Coast region. He 
attributed the absence of studies on these fossil echinoderms to neglect by 
paleontologists, not a poor fossil record. Much work on fossil echinoderms, 
particularly fossil echinoids, has been completed in the nearly 60 years since 
Howe wrote his paper. Unfortunately, neglect apparently continues to plague the 
comatulid crinoids. 

Oyen (1992) first reported the occurrence of H. bassleri in Florida from the 
Inglis Formation (Lower Ocala Limestone) in an abstract, and later published a 
detailed discussion of the biostratigraphic distribution of crinoids in Florida (Oyen, 
1995). The most recent discussion of Tertiary comatulids from Florida, as well as 
other states in the southeastern U.S.A., was by Oyen and Perreault (1997). All 
the Coastal Plain states from Louisiana to North Carolina have records of 
comatulids, and states such as Georgia and Alabama have several different taxa, 
some of which prove to be common when microscopic fractions are examined 
closely (Oyen and Perreault, 1997; Oyen, unpublished data). Therefore, I 
believe it is likely that more taxa will be found to augment my currently reported 
comatulid diversity for the state of Florida. 


279 


The fossil crinoids from Florida consist of several different skeletal 
components from two taxa of comatulids (Table 4-1). The collection of H. bassleri 
consists of skeletal components including 50 centrodorsals, 53 radial plates, and 
20 basal rays found at UF locality CI001 (UF 39054 - UF 39090) in Citrus 
County. Specimens are small, with most centrodorsals less than 10 mm in 
diameter, and the effects of diagenesis (recrystallized ossicles, epitaxial 
cements) are visible in many of the fossils. Figure 3-40, parts A-C show 
representative views of the dorsal and ventral surfaces of the centrodorsal, the 
included radial plates and basal rays for this species, as well as the imperfect 
preservation state of the crinoid components. 

A second comatulid crinoid species from Holmes County, Florida was 
collected from the Upper Ocala Limestone. Skeletal components consist of five 
brachial plates and only one centrodorsal from UF locality HO001 (UF 48125 and 
UF 48126). Figure 3-40, D-G, show dorsal and ventral views of the centrodorsal 
and two representatives of the five brachial plates. These specimens are 
distinctly smaller than those of H. bassleri , with the centrodorsal measuring 
approximately 2.0 mm in diameter and the brachials averaging 1 .0-1 .5 mm in 
length. The taxonomic status of these specimens is still uncertain. 

Asteroids 

The record of asteroid taxa from the Cenozoic of Florida is rather limited, 
even though the abundance of skeletal fragments may be high in some strata. 
Most echinoderms disarticulate soon after death, but the asteroids seem 


280 


particularly susceptible to leaving a complex fossil record for paleontologists to 
decipher. Individual sea stars may have hundreds of ossicles that disperse 
readily following death in higher energy environments. Only if burial occurs 
rapidly at the time of death or shortly thereafter will the original morphologic 
arrangement remain intact. Records of asteroids exist from the Paleozoic 
through the Cenozoic, but in general, asteroids are rare and any well-preserved 
fossils are exceptional (Spencer and Wright, 1966). 

Approximately 1,800 species of sea stars have been described from the 
world’s oceans, with the richest diversity in reefs from the Red Sea and Indo- 
West Pacific (Hendler et al., 1995). They also report that the northern Caribbean 
and Gulf of Mexico have relatively low diversity, with only 18 species in the 
shallow depths of less than 46 m and another 160 species at depths down to 
3658 m. Camp et al. (1998) compiled a list of shallow-water marine invertebrates 
from Florida’s coast and reported 21 species of asteroids from the depth range of 
0 to 37 m. The fossil record of asteroids from the Cenozoic of Florida is 
extremely limited in comparison with the modern faunal diversity. One of the 
reasons for the low fossil diversity simply is due to the difficulty in identifying 
lower taxonomic levels based on isolated ossicles. 

Fossil asteroids in Florida are known from the Middle Eocene through the 
Pleistocene. Only one group of specimens from the Pliocene has been identified 
to species level, one Pliocene sample has been identified to genus level, while 
the rest of the fossils currently have been limited to tentative generic or familial 


identifications. 


281 


The Paleogene has records of asteroids from the Middle Eocene Avon 
Park Formation and the Late Eocene Ocala Limestone. A beautiful fauna 
associated with fossil seagrasses in the Avon Park (Ivany et al., 1990) contains 
juvenile echinoderms, including sea stars representing possibly Goniodiscaster , 
a member of the family Oreasteridae (Figure 3-40, H). This same taxon is 
represented by several adult specimens, both as articulated calcite ossicles and 
external molds, and as numerous isolated marginal ossicles from several Late 
Eocene exposures (Figure 3-40, l-J; Figure 3-41, A-B). Figure 3-42 A shows an 
example of a Recent Florida Oreaster reticulatus (Linnaeus, 1816) for 
comparison. Unfortunately, specimens normally are too poorly preserved to 
permit a more confident taxonomic identification. In the Oligocene, asteroid 
ossicles have been reported from the Suwannee and Bridgeboro limestones. 
Once again, no complete specimens exist (i.e., predominantly isolated ossicles), 
so the best interpretation is that these samples are skeletal components from 
one or more species belonging to the Oreasteridae. 

Asteroid fossils and skeletal fragments are found throughout the Neogene 
as well. Several formations from the Miocene contain asteroid ossicles, all of 
which have been interpreted as possible representatives of the family 
Astropectinidae (see Figure 3-42, B-C for modern example). These ossicles are 
much smaller than the typical Paleogene sea star ossicles. Thus, they may be 
overlooked more easily, even by workers familiar with large marginal ossicles 
from the older rock units. Formations that have astropectinid ossicles include the 


282 


Arcadia, Coosawhatchie, Parachucia, and Chipola (where such ossicles are the 
most common). 

The most spectacular preservation of fossil sea stars in Florida is found in 
the Pliocene. Jones and Portell (1988) described the occurrence of whole body 
asteroids, Heliaster microbrachius Xantus, from the Tamiami Formation in 
Charlotte County, Florida. This species has between 27 and 44 arms, and many 
of the individuals (of the approximately 360 total) are preserved complete with 
arms intact (Figure 3-43 A). The Tamiami Formation has two other taxa as well, 
with articulated ossicles of partial arms of Luidia sp. (Figure 3-43 B) and ossicles 
identified as a possible Oreasteridae. Another Pliocene formation with sea star 
skeletal debris is the Caloosahatchee Formation, which contains ossicles that 
await full identification. Finally, two Pleistocene stratigraphic units (Bermont and 
Nashua formations) have asteroid ossicles that have yet to be specifically 
identified. 

Ophiuroids 

Ophiuroids have a very poor fossil record in the Cenozoic of Florida. Like 
the sea stars, these organisms have many individual skeletal plates that 
disarticulate readily after death. The ossicles typically are very small (most are 
less than one mm in diameter), and are difficult to find without careful 
examination of the sedimentary rock. Furthermore, such isolated ossicles may 
be overlooked because they are unfamiliar to many paleontologists. Thus, the 


283 


paucity of fossil ophiuroids probably is an artifact of the sampling habits of 
paleontologists to a greater degree than a real lack of fossils. 

The diversity of brittle stars in the modern oceans is estimated to be 
approximately 2,000 species (Hendler et al., 1995). Hendler et al. report 
ophiuroids associated with coral reefs in the Caribbean to be moderately 
abundant, and can be found in densities of 20 to 40 individuals per m^. 

Burrowing species within soft sediment can be concentrated up to 100 times as 
dense as the reef species. In the shallow marine environment of Florida today, 
there are 65 species of ophiuroids at depths less than 37 m (Camp et al., 1998). 
Therefore, it would be reasonable to expect a much better fossil record of these 
organisms than I have found in Cenozoic strata of Florida. 

There are only three formations with records of ophiuroids in the state. 
Ivany et al. (1990) reported complete external molds of juveniles found along the 
surface of fossil seagrass blades in the Eocene Avon Park Formation (Figure 3- 
40 H), but no identification of specific taxa was included in the paper. Ophiuroid 
vertebral ossicles also are found in the Miocene Marks Flead Formation, but no 
lower taxonomic identification has been made for these fossils (Figure 3-43 C). 
Finally, a single complete specimen (Figure 3-43 D) associated with the asteroid 
Heliaster microbrachius is present in the Pliocene Tamiami Formation in 
southwestern Florida, but no specific taxonomic identification has been 
determined due to the specimen’s recrystallization and imperfect preservation. 


284 


Biases in the Cenozoic Echinoderm Record 

The pattern of echinoid diversity from the Middle Eocene through the 
Pleistocene is influenced by several variables. At this time, my echinoid data are 
much more detailed and thorough than the asteroid and ophiuroid data (both 
stratigraphic and taxonomic). Therefore, most of the following discussion will 
focus on the echinoids. 

As noted in the taxonomic section, the echinoid diversity pattern now 
known from Florida has changed from previously published information. In 
particular, the diversity record in the Miocene has improved dramatically. In this 
section I will address how biases may be influencing the diversity pattern for 
echinoderms in Florida and possible reasons why the earlier data were relatively 
sparse for the Neogene (especially in the Miocene). 

Resolution of Data 

The data presented herein are at a relatively coarse level with respect to 
time units (epoch level resolution) and stratigraphic units (formation level 
resolution). Data have been gathered from publications, museum collections, 
and amateur and professional collectors. In some cases only locality information 
was provided, and depending on my knowledge of the locality and any 
associated quarries at the locality, I can determine stratigraphic unit(s) 
associated with the locality. I have been very conservative and cautious 
regarding these data derived from outside my personal work, and any 


285 

questionable stratigraphic assignments have been omitted from this database 
until better stratigraphic control can be established. 

A second stratigraphic-control problem that limits resolution of data to 
formations only is that samples may have been collected from spoil piles. 
Therefore, exact stratigraphic position cannot be determined for those specimens 
and only the formation can be assigned to these specimens. Knowledge of the 
pit depth, mining depth for specific spoil piles, observations of sections during 
excavations, and discussions with quarry operators all are taken into 
consideration before assigning stratigraphic information to echinoid samples 
used in this study. Relatively few published records of echinoids and their 
stratigraphic position exist; therefore, the resolution for general patterns is limited 
to presence or absence in specific formations. 

Stratigraphic Nomenclature 

One of the greatest difficulties in geologic work in Florida involves 
distinguishing unique stratigraphic units according to the guidelines of the North 
American Stratigraphic Code. In many cases, only subtle variations in lithology 
exist among the state’s Cenozoic strata (in contrast to non-Coastal Plain areas), 
which makes defining formations a challenge. A second inherent problem in the 
stratigraphic nomenclature of Florida has been the reliance on fossils to define 
formations. Examination of the literature involving names and definitions for 
several Cenozoic units shows that faunal constituents were the primary basis for 
identifying these formations. Unfortunately, much of the original work was done 


286 


prior to development of formal stratigraphic codes or guidelines and these 
formations have become strongly entrenched in the literature. Examples include 
the formations of the Eocene Ocala Group (i.e., Inglis, Williston, and Crystal 
River formations), which are now all part of the Ocala Limestone, and some 
Pliocene units such as the Caloosahatchee and Nashua formations. This type of 
nomenclature influence could be present in varying degrees from the Eocene 
through the Pleistocene. Thus, readers must be aware that such information is, 
in part, a function of the interpretations of lithostratigraphers, sedimentologists, 
biostratigraphers, and paleontologists to find geologically sound and functional 
solutions to the stratigraphic nomenclature difficulties in the state. 

Outcrop Exposure and Relief 

Another bias in the echinoderm data is a function of outcrop exposure. It 
is obvious that formations which have only limited surface exposure will have 
less geologic and paleontologic data readily available to gather during fieldwork. 
In Florida, only rocks from the Middle Eocene through the Pleistocene are 
exposed at the surface, and rocks from each epoch are represented by unequal 
proportions of surface exposure. This means that some epochs are less likely to 
be sampled. Furthermore, even when exposed, accessibility to some outcrops 
can be extremely difficult. For example, in the panhandle region where the 
Intracoastal Formation outcrops, very few roads exist, and those that do consist 
of narrow, dirt logging roads through rough, undeveloped areas. 


287 


A second aspect of collecting availability is the dependence on quarrying 
operations. Several mines have provided access for paleontologists and 
collectors. As a result, some stratigraphic units that may not be exposed directly 
on the surface may be exposed in the shallow subsurface following mine 
excavations. This allows fossils to be collected from areas where a given 
formation may not be represented on a geologic map for the area. 

Perhaps just as important is the bias associated with the collectors of 
fossils themselves. Most collectors fall into two categories: “fossil shellers” or 
“bone hunters”. Therefore, these collector groups tend to have a relatively 
narrow range of taxonomic focus. Shellers focus primarily on the mollusks while 
the bone hunters look for vertebrate remains. Other taxonomic groups such as 
the echinoderms, arthropods, and cnidaria have had a relatively low priority 
among these collectors, and therefore fewer specimens have been added to 
museum collections. This trend seems to be changing (specifically, it is 
improving) as more collectors are informed of the need for such fossil taxa by 
museums and paleontologists. 

One of the rock exposure artifacts that also may help explain the large 
increase in Neogene taxonomic and stratigraphic records is the increasing 
population growth in Florida. As development and construction occur in the fast- 
growing areas along the coasts and in south Florida, more land is excavated for 
buildings, highways, utilities, and other infrastructure needs. These excavations 
provide opportunities to examine sedimentary rocks that simply were unavailable 
prior to development activities. On occasion, these temporary pits have exposed 


288 

units containing rare fossils such as the sea stars, Heliaster microbrachius 
(Figure 3-41). 

Finally, another factor in contributing to bias regarding exposure of units is 
the general topography of the state. Florida has limited relief, with the highest 
point in the state just 104 m above sea level (Schmidt, 1997). South Florida, in 
particular, has a maximum topographic variation of less than about 7.5 m, and 
averages only a few meters locally. Strata tend to be relatively flat-lying and, 
therefore, only the top stratigraphic unit will be exposed unless excavation 
beneath the surface unit is completed. In addition, many areas of south Florida 
are covered with heavy vegetation, swamps, and marshes, which further reduce 
outcrop exposure in the area. Lack of surface outcrops is not limited to south 
Florida, however, and even though relief may be greater in the northern 
peninsula and panhandle areas it still is limited in comparison to non-Coastal 
Plain terrain. Overall, the Neogene units (found in lower relief areas) appear 
more likely to have suffered from lack of outcrop exposure than the Paleogene 
units due to these factors. 

Carbonate Versus Siliciclastic Environments 

Yet another bias in the echinoderm record may be related to the 
preservation potential for fossils in carbonate versus those in siliciclastic facies. 

My study has no experimental evidence to quantify this proposal, but personal 
observations of fossils in outcrop and in museum collections generally show that 
less abrasion and fragmentation of specimens is present in the carbonate rocks 


289 


in contrast to the fossils found in siliciclastic sedimentary rocks. Florida’s non- 
carbonate rocks are dominated by quartz sands, with lesser amounts of 
phosphate grains, heavy minerals (such as ilmenite or rutile), and clay minerals. 
Does the high content of abrasive minerals (i.e., quartz) have a significant impact 
on the quality of preservation in fossils? Are softer and less durable carbonate 
minerals (i.e., calcite and aragonite) less likely to create deleterious effects on 
the skeletal components within the sediment that produces a sedimentary rock? 

It seems logical those questions can be answered affirmatively, but the answers 
probably are not so simple. 

Chave (1964) published one of the first papers on skeletal durability and 
preservation potential, in which he tumbled skeletal material in a rock tumbling 
barrel using various abrasives to provide durability estimates. Chave’s study 
used a variety of taxa for the experiment, including the regular echinoid 
Stronqylocentrotus and the sea star Pisaster . as well as mollusks, coral, algae, 
and bryozoans. Chave varied the type of sediment included in the tumblers with 
the skeletal material, among chert pebbles, sand grains, and skeletal parts only. 
His results showed that the siliceous chert and sand particles produced subequal 
abrasion and degradation results, and both produced more effective degradation 
than when only the carbonate shell materials were impacting one another in the 
tumblers. One of the more important conclusions of his study, however, was not 
that siliceous grains are more detrimental to fossil preservation than skeletal 
grains, but, instead, skeletal microarchitecture is the most critical factor in 
determining the rate of degradation. Unfortunately, this does not answer my 


290 


questions regarding physical degradation and its relationship to sediment 
composition since Chave’s experiment was not run using carbonate sediment as 
an abrasive. 


Taphonomy of echinoderms has been discussed by a number of workers 
in recent years, including Schafer (1972), Kidwell and Baumiller (1990), Allison 
(1990), Donovan (1991), and Greenstein (1991, 1992, 1993). Detailed 
discussions of preservation and processes are included in these works, but none 
has specifically addressed differences in carbonate and clastic environments with 
respect to echinoderm preservation potential. 

Research on foraminiferal taphonomy by Kotler et al. (1992) integrated 
both physical and chemical tests to determine preservation styles and processes 
for the microorganisms. Biogenic carbonate sands were used as the primary 
abrasive agent while quartz sands were used only as a matrix to identify post- 
mortem transport of the forams. This work produced slightly different results 
compared with abrasion tests done by other workers using siliciclastic sediment. 
For example, Chave (1964) found skeletal architecture to be important in 
differentiating exposure effects among selected macroinvertebrate taxa, whereas 
Kotler et al. (1992) found only limited variation in abrasion resistance among the 
microinvertebrate foraminifera that may be attributed to skeletal architecture. 

Their results showed that other factors including test hydraulic behavior, low 
traction velocity, and test shape, have discernable effects on abrasion 
susceptibility. Comparative observations of Florida echinoid fossils from the 
Paleogene and Neogene units suggest that abrasion is not the most important 


291 


taphonomic variable of concern. Instead, the overall preservation state of the 
echinoids, which includes the extent of fragmentation, chemical alteration 
(particularly dissolution), as well as abrasion, is better for the carbonate- 
dominated Paleogene formations, and poorer in siliciclastic-dominated Neogene 
formations. 

Several reasons why carbonate and siliciclastic environments produce 
different fossil preservation styles were reviewed in Kotler et al. (1992), and may 
be applicable to Florida material. Carbonate beds or shell-rich clastic beds tend 
to produce a favorable chemical environment by buffering pore fluid against 
dissolution processes (Kidwell, 1989), while siliciclastic units with minimal 
carbonate shell content are more susceptible to dissolution. Rates of dissolution 
in terrigenous environments also are influenced by sedimentation rate, shell 
input, organic matter, and bioturbation (Meldahl, 1987). Furthermore, rate and 
consistency of sedimentation differ between carbonate and siliciclastic 
(terrigenous) environments, with long-term carbonate deposition typically slower 
than long-term terrigenous clastic deposition (Schindel, 1980). A combination of 
these factors would influence the preservation potential of calcareous shells, and 
the resulting differential preservation produced in carbonate versus siliciclastic 
regimes in turn would affect the diagenetic potential (i.e., alteration) that the fossil 
assemblage would undergo after burial (Kotler et al., 1992). 

Details regarding skeletal breakdown in carbonate versus siliciclastic 
environments are extremely limited; thus, I may only speculate on the 
depositional environment’s influence on the echinoderm record of Florida. 


292 


However, such a pattern is observable in my data, with the Neogene units 
showing distinctly larger increases in taxonomic and stratigraphic records in 
contrast to Paleogene units. Therefore, I believe this has contributed to a 
diversity bias, which I am addressing in this work. 

Age of Stratigraphic Units 

Age determination of stratigraphic units also will affect the echinoderm 
diversity pattern for the Cenozoic of Florida. I have not attempted to present 
information regarding absolute ages for the formations discussed in this study, 
and there are significant uncertainties. The Caloosahatchee and Nashua 
formations, for example, are mapped as Plio-Pleistocene in age but I have 
treated the echinoids found within these formations as being Pliocene. In reality, 
these formations are Plio-Pleistocene units, with the upper beds of the units 
Pleistocene in age. I do not believe that these formations are restricted to only 
one epoch, but have chosen the single epoch which best represents the age of 
the echinoids found within those formations. Clearly, taxonomic diversity per 
epoch will change as age assignments of formations change, since my data are 
constrained by formation-level resolution. 

Epoch Duration 

One aspect of the Florida echinoid diversity pattern that is important to 
note is the influence of time variation for my epoch resolution data. My data are 
limited to epoch assignment, where each epoch is unequal in duration. 


293 


Therefore, even though the number of known taxa from each epoch varies from a 
minimum of six species in the Pleistocene to a maximum of 38 species in the 
Eocene, these data can be standardized to help minimize the temporal duration 
effect. Kier (1977), McKinney et al. (1992), and Donovan (1993) standardized 
echinoid diversity values by dividing the diversity by the epoch duration (in 
millions of years), and herein I follow this style as well. 

The Avon Park Formation is the oldest unit at the surface in Florida and 
the oldest unit containing echinoids. The age of the Avon Park is late Middle 
Eocene (Puri and Vernon, 1964), which I assume to represent the base of the 
Bartonian Stage at approximately 42.1 Ma (Harland et al., 1990). Using epoch 
boundary ages provided in Flarland et al., the duration of each of the epochs was 
calculated and used to determine diversity values normalized by millions of years 
of duration. Since the age of the Avon Park is only an estimate, the normalized 
diversity may be slightly skewed, but it is at least a reasonable estimate to use 
for comparative purposes. The results of the calculations are provided in Table 
4-6. 

Following time normalization, the pattern of echinoid diversity from the late 
Middle Eocene through Pleistocene changes noticeably in two areas. First, the 
Miocene diversity no longer is above average, but instead is nearly the lowest of 
the intervals examined. Second, the Pleistocene diversity changes from the 
lowest diversity interval to one of the highest diversity intervals (due to the very 
short duration of the epoch). As a result, the normalized Florida data (Figure 4-3) 
do not match normalized worldwide echinoid data because the Miocene diversity 


294 


Table 4-6. Normalized Florida echinoid diversity. Total echinoid diversity for 
each epoch is divided by the duration of the epoch to produce 
normalized diversity values (in number of species per million years 
[Ma]). Epoch duration values v\/ere calculated from data in Harland et 
al. (1990). 


EPOCH 

EPOCH LENGTH 
(Ma) 

ECHINOID TAXA 
TOTAL 

NORMALIZED 

DIVERSITY 

Pleistocene 

1.63 

6 

3.68 

Pliocene 

3.56 

23 

6.46 

Miocene 

18.10 

22 

1.22 

Oligocene 

12.10 

11 

0.91 

Eocene 

6.70 

38 

5.67 


NORMALIZED FLORIDA ECHINOID DIVERSITY 


X 

o 

o 

Q_ 

LU 


Pleistocene 


Pliocene 


Miocene 


Oligocene 


Eocene 



0.00 1.00 2.00 3.00 4.00 5.00 6.00 

NORMALIZED DIVERSITY (spp/Ma) 


7.00 


Figure 4-3. Normalized Florida echinoid diversity for species from the Middle 

Eocene through Pleistocene. Note the proportionally low normalized 
diversity for the Miocene and proportionally high diversity for the 
Pleistocene. This is in obvious contrast to the raw diversity pattern 
illustrated in Figure 4-1 . 


295 


of Florida is proportionally low whereas the worldwide diversity is proportionally 
high (Figure 4-4 A). Alternatively, the Florida diversity closely follows the pattern 
found in the Cenozoic of Jamaica (Figure 4-4 B). 

Donovan’s most recent diversity data for Jamaica (Donovan, in press), 
which includes both published and unpublished sources, has a slightly higher 
normalized diversity for the Oligocene than my records show in Florida. 

Flowever, the general pattern of lower than average diversities for the Late 
Paleogene (Oligocene) and Early Neogene (Miocene) is found in both regions of 
the Caribbean. The correspondence of Jamaica and Florida diversities likely 
suggests a regional divergence from the worldwide Miocene diversity increase, 
which may be due to sustained unfavorable environmental conditions in the 
Caribbean following the Eocene epoch. 

Taxonomic Nomenclature 

Taxonomic descriptions of fossil echinoderms in Florida are predominantly 
at the specific level, with few subspecies recognized. For the echinoids, 
published data show less than ten subspecies have been reported from the 
Cenozoic. I have taken a conservative approach for this study and omitted all 
subspecies from my “taxonomic records” count for all epochs. I believe closer 
examination of these taxa will show most or all such subspecies to be 
taxonomically invalid. Therefore, inclusion of subspecies only results in the 
inflation of taxonomic diversity for the Eocene, Pliocene, and Pleistocene epochs 
where such taxa are found. An example of possible echinoid taxonomic splitting 


296 


Myr BP 



0 20 40 60 80 

SPECIES Myf ' 


A) 


Myr BP 



B) 

Figure 4-4. Comparison of normalized diversity values. A) Worldwide 

normalized diversity of fossil echinoid species in the Cenozoic using 
data from Kier (1977). B) Normalized diversity of Cenozoic species 
from Jamaica. (Modified from McKinney et al., 1992) 


297 


is that of the clypeasteroids in the Pleistocene Bermont Formation, CIvpeaster 
rosaceus (Linnaeus) and CIvpeaster rosaceus dalli (Twitchell). This dissertation 
is not designed to formally revise echinoderm taxonomy, but it should be noted 
that the number of taxonomic records included in my values for the Pleistocene 
has only one record for C. rosaceus from the Bermont Formation rather than two. 
Additional research is continuing on taxonomy of Florida echinoids (by C. Oyen 
and R. Portell), crinoids (by C. Oyen), and asteroids and ophiuroids (by D. Jones 
and Portell), and the validity of various subspecific assignments will be 
considered in future work. 

A second taxonomic problem for biostratigraphy of echinoids in the 
southeastern U.S.A., is the geographic effect on taxonomy. Similar echinoids 
from different geographic areas have been called different species, even though 
the specimens are not sufficiently distinct to justify a split. For example, Arbacia 
crenulata Kier from the Tamiami Formation in Florida and A. improcera (Conrad) 
of the Yorktown Formation of Virginia and North Carolina, as well as the Croatan 
Formation of North Carolina, are found in different geographic areas, and 
therefore were described as different species. It has been noted subsequently 
that splitting these age-equivalent taxa into unique species is not valid, since 
further examination of specimens shows no significant morphological variation 
that would allow defining two distinct species (Kier, 1972). Fortunately for my 
work in Florida, such geographic-based taxonomic splitting has little to no impact 
on the Florida echinoid diversity because only one or two species, of those which 
may have this effect, were counted in my taxonomic records. 


298 


Collector Bias 

One of the biases of the fossil record that affects most taxonomic groups, 
including the echinoderms, is the tendency for people to collect clean, unbroken, 
and larger sized (that is, more obvious) fossils. Clearly, not all researchers have 
this style of sampling bias, but it often requires conscious effort to include 
incomplete specimens, small fragments, moldic samples, or bags of matrix for 
microfossil analyses. While it becomes more difficult to identify lower taxonomic 
levels from isolated skeletal components, such fragments can at least provide 
some data for paleoecology interpretations. This has been noted in echinoid 
taphonomy and paleoecology research by several workers, including Gordon and 
Donovan (1992), Greenstein (1993), Kidwell and Baumiller (1990), and Nebelsick 
(1992). 

The Miocene epoch has the greatest increase in stratigraphic and 
taxonomic records of echinoids in Florida. One of the reasons this is true is 
because it is not common to find complete, well-preserved echinoids in Miocene 
strata. Several of the new records I include in this paper are derived from test 
fragments, spines, or molds, rather than nearly complete tests. An example of 
the moldic preservation style is found in Lovenia clarki from the Chattahoochee 
Formation (Figure 3-18), and silica rubber peels are often made of the mold in 
order to more detailed descriptive work. In some stratigraphic units these 
fragments are not uncommon, but probably were ignored because of the difficulty 
in sorting and identifying such skeletal parts to family, genus, or species levels. 
Furthermore, some of the fragments are relatively small and may be found only if 


299 


one examines the fine fractions of sediment in section or in spoil piles. This bias 
is not limited to the Miocene units but seems to be proportionally less prevalent in 
other portions of the Cenozoic of Florida. 

One excellent example of ho\A/ to avoid such biases by careful 
examination and collection of microfossils and incomplete specimens is that of 
Harold and Emily Yokes (Tulane University). When collecting the numerous 
Chipola Formation localities in northern Florida (along Tenmile Creek, Farley 
Creek, and the Chipola River), they and their associates would not only collect 
the taxonomic groups they were interested in personally, but also anything else 
of either micro- or macro-size. It is because of their comprehensive collecting, 
washing and sorting, that I can confidently make statements about echinoderms 
like Prionocidaris being a common and abundant fossil occurring in the Chipola 
Formation, even though the Yokes’ focused their research on mollusks rather 
than echinoderms. 

Substrate and Facies Preferences of Echinoids 

Echinoids are now known from nearly all habitats ranging from intertidal to 
deep marine, and from tropical to polar regions in all oceans throughout the 
world. Although a few species have nearly cosmopolitan distribution, most are 
geographically restricted and, even further, are habitat limited in distribution 
(Smith, 1984). Ernst Mayr (1954) described numerous ways the geographic 
distribution of echinoids is controlled. Mayr believed the most influential controls 
of echinoid species global distribution to be the presence of land barriers, wide 


300 


ocean basins, and oceanic current patterns. However, even within small 
geographic regions, species distribution is patchy, rather than continuous (Smith, 
1984). This non-continuous distribution is easily observed in the fossil record of 
echinoids in Florida strata, and thereby warrants a brief discussion of this pattern. 

In general, there are multiple controlling factors for the irregular 
distribution of echinoid species. The studies by Kier and Grant (1965), Ebert 
(1971), and Smith (1984) have defined eight factors. These controls include the 
substratum, hydrodynamic regime, predation, salinity, temperature, food 
availability, water depth, species behavior, and chance. Of these factors, the 
nature of the substrate (specifically, the physical characteristics including the 
grain size, sediment stability, degree of sorting, content of organic matter, 
porosity, and permeability) is most influential on local distributions (Smith, 1984). 

Sedimentary facies are defined by the physical characteristics of the 
sediment that indicate environmental conditions at the time of deposition. 
Individual species of echinoids have well-defined preferences for specific 
substrate characteristics; therefore, the presence of a given echinoid species 
may serve as a proxy indicator of a sedimentary facies for those paleontologists 
who have not examined the sedimentary rock that contained the fossils. 

Zoologists studying modern taxa have described the close relationship of taxa 
with specific substrate characteristics. One example is the research completed 
by McNulty et al. (1962) in which they examined the distribution of echinoids in 
the shallow water region of southern Florida. They found the spatangoid Moira 
atropos (a species first reported as a fossil from Florida in this dissertation) was 


301 


found only in sediments consisting of 75-90% sand-sized particles with 0-12% 
gravel and 10-20% silt, while mellitid species live only in medium- to fine-grained 
sands. Similar correlations between modern species and substrate 
characteristics have been reported for both regular and irregular echinoids by 
Lawrence and Ferber (1971), Heatfield (1965), and Smith (1980). Such a 
concept may prove to be a useful tool for sedimentary petrologists, 
stratigraphers, and paleontologists alike, because museum collections of 
echinoids that have associated stratigraphic and geographic information 
associated with their fossil echinoids can then be used to produce general facies 
distribution maps. I have included examples of such interpretations for selected 
Florida species in subsequent text and figures. 

Analyses of substrate preferences for a variety of Paleogene echinoids 
from the southeastern U.S. and other regions worldwide have been completed in 
the last dozen years. Burchard Carter of Georgia Southwestern State University 
has lead several such projects that have included fossils collected from Florida or 
species in nearby states that are also part of the fossil record in Florida (see 
Carter, 1989; Carter etal., 1989; Carter, 1990; Carter and McKinney, 1992; 
Carter, 1997). As part of his research. Carter produced thin-sections of both the 
limestones and the echinoid tests (with sediment still trapped within the internal 
cavity of the test) for determination of grain size of the substrate. He found 
consistent patterns among the irregular echinoids that allowed him to identify 
substrate preferences for specific and/or generic levels of taxa. 


302 


Two of the publications that resulted from this work, Carter et al. (1989) 
and Carter (1990), indicated species that were identified as dominantly inhabiting 
one (or rarely, two) of the three qualitative categories of carbonate substrates 
termed clean sand, muddy sand, or very muddy sand. As an example, all three 
species of Oligoovgus (O. ohelani . O. haldemani . and O. wetherbyi ) were 
determined to be abundant or common in clean sand substrates (see figure 1 in 
Carter, 1990, for a complete listing of taxa and substrate preferences). All 
Eocene species of Rhvncholampas also fall into the clean sand dweller category, 
while other species such as Eupataqus ocalanus and Wvthella eldridqei are 
muddy sand dwellers. Those species that occupy the very muddy sand category 
include Brissopsis steinhatchee and Eurhodia patelliformis , among others. 

These results are used as the primary guide for my projections of facies 
distributions via echinoid presence, although my own observations of facies 
preferences by particular echinoid species also correlate well with the data 
gathered by Carter and his co-workers. The work completed by Carter and 
others is limited, however, to the Late Eocene formations of Florida (using the 
Ocala Group stratigraphic nomenclature rather than the Ocala Limestone). 
Therefore, I have extrapolated those interpretations to include closely related 
echinoid taxa for the rest of the Cenozoic epochs based upon my field 
observations of both fossil and modern species from Florida and the shallow 
marine coastal areas. 

To illustrate the use of echinoids as sedimentary facies proxies, 1 have 
identified at least one or two species from the Eocene through Pleistocene 


303 


epochs to use as sand facies indicators. These species then are used to 
graphically display areal distribution of sand facies units within a given formation 
in the state of Florida. One limitation of this approach using my data should be 
noted, however. All data are limited to county level resolution for my areal maps. 
In reality, sand facies within a given formation often are far more limited, both 
laterally and stratigraphically, than would be indicated by my maps. The goal of 
these illustrations is to show the future value of using the echinoids as a tool, but 
recognizing further data must be gathered to fully accomplish this goal. As noted 
in earlier chapters, much of the stratigraphic data for the echinoids are limited to 
formation level only. Until stratigraphic sections are measured and specific 
geographic data such as latitude and longitude are included, these 
interpretations should be regarded as only a first attempt to apply the echinoid 
distribution data to other geologic research, such as sedimentary geology. The 
echinoids chosen to illustrate sand facies distribution throughout the state over 
time include: 1 ) Oliqopyqus wetherbvi and Rhvncholampas ericsoni for the Ocala 
Limestone of the Eocene (Figure 4-5); 2) Rhvncholampas qouldii for the 
Suwannee Limestone of the Oligocene (Figure 4-6); 3) Abertella aberti for the 
Arcadia Formation of the Miocene (Figure 4-6); 4) Encope tamiamiensis for the 
Tamiami Formation and Rhvncholampas avresi for the Caloosahatchee 
Formation of the Pliocene (Figure 4-7); and 5) CIvpeaster rosaceous for the 
Bermont Formation and Mellita quinquiesperforata for the Satilla Formation of the 
Pleistocene (Figure 4-8). A common theme in these taxa is that all are members 
of one of the orders of cassiduloids, oligopygoids, or clypeasteroids. Species of 


304 




Figure 4-5. Eocene echinoids as carbonate sand facies distribution indicators, 
(a) Areal distribution of carbonate sand facies in the Ocala 
Limestone as indicated by Oliqopyqus wetherbvi . (b) Areal 
distribution of carbonate sand facies in the Ocala Limestone as 
indicated by Rhvncholampas ericsoni . 



305 



Figure 4-6. Echinoids as carbonate sand facies distribution indicators, (a) Areal 
distribution of carbonate sand facies in the Oligocene Suwannee 
Limestone as indicated by Rhvncholampas qouldii . (b) Areal 
distribution of carbonate sand facies in the Miocene Arcadia 
Formation as indicated by Abertella aberti . 



306 




Figure 4-7. Pliocene echinoids as sand facies distribution indicators, (a) Areal 
distribution of sand facies in the Tamiami Formation as indicated by 
Encope tamiamiensis . (b) Areal distribution of sand facies in the 
Caloosahatchee Formation as indicated by Rhvncholampas avresi . 




307 



Figure 4-8. Pleistocene echinoids as sand facies distribution indicators, (a) 
Areal distribution of sand facies in the Anastasia and Satilla 
formations as indicated by Mellita quinquiesperforata . (b) Areal 
distribution of sand facies in the Bermont Formation as indicated by 
CIvpeaster rosaceous . 




308 

these orders, in turn, share similar habitat preferences, only one of which is 
substrate characteristics. In some circumstances, these fossils are not 
exceptionally abundant within each of these formations, but they are at least not 
rare within the units and thereby would allow a reasonable likelihood of recovery 
during fieldwork so they may be applied to facies distribution interpretations. 

In summary, the substrate preference of given species of echinoids will 
not only influence their geographic distribution throughout the state, but the 
presence or absence of a given facies will also ultimately control the abundance 
and potential diversity of echinoids during a given time. Therefore, the diversity 
pattern present from the middle Eocene through the Pleistocene is influenced in 
part by the depositional environment and facies that existed at any given time. 

The Eocene-Oligocene Diversity Change 
Several significant changes occurred in the world's oceans in the middle 
Paleogene which are readily identified based on sedimentation patterns, isotopic 
studies of various fossil taxa, and paleobiogeographic patterns. In particular, 
paleobiogeographic patterns reflect distribution changes for fossil taxa that, in 
turn, reflect the change in environmental conditions present in the water masses. 
Extinction of various taxonomic groups also is associated with the Eocene- 
Oligocene boundary and was extensive enough to be noted slightly above 
background extinction rates. However, it was not as pervasive as extinction 
levels at other times such as the Permian-Triassic and Cretaceous-Tertiary 
boundary events (see Raup and Sepkoski, 1982 and 1986). 


309 


The extinction event at the Eocene-Oligocene boundary was called the 
"terminal Eocene event" by Wolfe (1978), who noted significant changes in 
terrestrial plant biogeographic distributions as well as in global temperature 
patterns. Temperature change (cooling) in the late Eocene has been noted in 
marine taxa both by faunal assemblage shifts from warm to cool water taxa 
dominance as well as isotopic shifts in ^®0. Specifically, these changes can be 
seen in benthic foraminifera (Buchardt, 1978; Corliss, 1979; Keigwin, 1980; 
Keigwin and Corliss, 1986), planktonic foraminifera (Berggren, 1977; Keller, 1983 
and 1986; Boersma, 1987), ostracodes (Benson et al., 1984), molluscs (Hansen, 
1987), and echinoids (McKinney and Oyen, 1989; McKinney et al., 1992). 
Evidence for dramatic climatic change (specifically, temperature drop) is also 
supported by extinction of terrestrial vertebrate groups at the end of the Eocene 
(Prothero, 1985). 

Changes in various marine invertebrate taxa as well as the proposed 
forces for the change are discussed in the following paragraphs. The influence 
of temperature change, tectonic activity, development of the psychrosphere, sea 
level change, and high latitude glacial buildups are all important interrelated 
factors that had an effect on faunal shifts and extinctions in the marine realm 
near the end of the Eocene. Thus, the significant diversity drop in Florida 
echinoids (shown earlier in this chapter) may be controlled by one or more of 
these changes taking place in the Paleogene. 


310 


Early Paleogene Oceanographic Conditions 

The early to middle Paleogene oceans represent a transition from 
thermospheric circulation to predominately thermohaline circulation as latitudinal 
temperature gradients gradually increased (Haq, 1981). This thermal gradient is 
a function of increased cooling in high latitude regions while low latitudes 
maintained a reasonably stable temperature regime. Although the forces driving 
circulation were beginning to change, during the Paleocene the world's oceans 
were still dominated by patterns established in the Cretaceous. These include 
the paleo-Gulf Stream flowing northward and the continuous equatorial current of 
the Tethys Seaway that dominated tropical circulation with westward flow (Haq, 
1981). This represented the major flow path for inter-oceanic circulation 
connecting the Indo-Pacific and Atlantic-Pacific oceans (Kennett, 1982). 

The thermospheric circulation (Cretaceous) that established the currents 
for the early Paleogene was based on a system without as strong of a thermal 
gradient between equatorial and polar latitudes. Further, the vertical thermal 
gradient in the oceans was less at the beginning of the Paleogene (Haq, 1981), 
thereby suggesting that both surface and deep water circulation was sluggish in 
comparison with the late Paleocene to early Eocene thermohaline circulation. 

Land mass arrangement was much different at the beginning of the 
Paleocene than at the end of the Eocene, and this played a critical part in 
circulation and temperature/climate patterns. Antarctica, South America, and 
Australia were still joined and concentrated in the Southern Ocean over the polar 


region, but this arrangement had been altered tectonically by the late Eocene 
and represents a critical event in Cenozoic paleoceanography (Kennett, 1977). 

Early To Middle Paleogene Transitions 

Kennett (1977) described several important changes in the 
paleoceanographic characteristics during the Cenozoic. These major changes 
influenced not only the climatic traits of the earth but also affected the biota 
associated with the oceans. Changes include: 1.) Separation of Australia and 
South America from Antarctica allowing development of the circum-Antarctic 
Current, 2.) Disturbance of the equatorial Tethyan Seaway, and 3.) Development 
and interaction of high latitude bottom waters as a result of climatic and glacial 
events in those regions. 

At the beginning of the Paleocene, the Antarctic continent (including still- 
attached Australia) was located in a polar position with no significant glacial ice 
coverage and both shallow and deep waters were relatively warm (Kennett, 

1982). The north polar latitude Arctic Ocean was closed in both by the Atlantic 
and Pacific Oceans, thereby closing off cold bottom water connections in the 
northern high latitudes. Kennett (1982) notes that this partial isolation of bottom 
waters caused geochemical variations in the world oceans such as the carbonate 
compensation depth (CCD) rising through the early and middle Paleogene in the 
Pacific, while lowering during the same time in the Atlantic. Haq (1981) 
suggested the Paleogene tectonic conditions reduced the northern area in the 
Pacific, resulting in an increased importance of the southern high latitudes as a 


311 


312 


source area for deep waters for the Indian and Atlantic Oceans in the middle and 
late Tertiary. 

Haq (1981) cited several important changes through the Paleogene in 
addition to those given by Kennett (1 977). Prior to the beginning of separation of 
Antarctica and Australia, separation of Greenland and Scandinavia began in the 
northern hemisphere, which in turn initiated the formation of the Greenland- 
Norwegian Sea. This did not significantly affect circulation patterns in the oceans 
during the early Paleogene, but by the middle Paleogene a surface water 
exchange with the Atlantic was established. The Greenland-lceland-Faroe Ridge 
system that had previously blocked water exchange is believed to have subsided 
enough by the latest Eocene to exchange cold, dense water into the Atlantic 
(Haq, 1981). However, significant outflow of North Atlantic Deep Water (NADW) 
was not achieved until after the Paleogene, when subsidence of the Greenland- 
lceland-Faroe Ridge had progressed further. 

The late Eocene also was a time in which Tethyan circulation was 
beginning to be restricted somewhat in the northern Indian Ocean as India 
migrated toward Asia (Kennett, 1982). This was not a dramatic constraint on the 
current flow however, and likely only represents a minor oceanographic change 
at the end of the Paleogene. 

Perhaps the most important oceanographic change to take place in the 
Paleogene is the tectonic separation of Australia from Antarctica. During the 
Paleocene, the Australia-Antarctica continent was in a polar position (Kennett, 
1977) but during the early to middle Eocene, Australia began to migrate 


313 


northward creating an ocean between the two continents (Haq, 1981; Kennett, 
1982). The impact of this separation was not realized in the world's oceans 
immediately because only very restricted circulation surrounding Antarctica was 
permitted due to the presence of the South Tasman Rise and the not yet open 
Drake Passage. 

By the late Eocene (~40 Ma), a shallow water connection between the 
Indian and Pacific Oceans had been established after the South Tasman Rise 
had subsided (Kennett, 1977; Haq, 1981). This is an influential factor in the 
circulation patterns of the oceans because it initiated the establishment of the 
circum-Antarctic Current. Although the circum-Antarctic Current was not fully 
developed until the opening of the Drake Passage was complete sometime in the 
Oligocene (-38-30 Ma [Kennett, 1977 and 1982; Haq, 1981]), its influence was 
felt as early as 40 Ma. 

The greatest effect of the isolation of Antarctica was the thermal change in 
ocean waters, initially cooling the surface waters and later causing a drop in 
bottom water temperatures. Kennett (1982) describes surface water temperature 
drops of 10° C from the early Eocene (at 20° C) to the late Eocene (at 10° C). 

Haq (1981) noted that this cooling was significant enough to cause large-scale 
freezing at sea level near Antarctica, and by the end of the Eocene (38 Ma), the 
initiation of production of Antarctic Bottom Water (AABW). 

Another important feature associated with the cooling of surface and, in 
particular, bottom waters is the development of the pyschrosphere and the 
modern two-layered ocean. Benson (1975) first described this event as 


314 


occurring at approxirnatGly 40 Ma bas6d on a cold bottom Iay6r psychrosphoric 
benthic ostracode fauna. Therefore, this marks the beginning of thermohaline 
dominant circulation as is present in the modern oceans (Kennett, 1977), versus 
thermospheric circulation as was present in the Cretaceous and much of the 
Paleogene. It is important to note that while many workers would describe the 
terminal Eocene event as the most significant paleoceanographic change in the 
Cenozoic (Kennett, 1982), the biotic crises may not actually reflect event(s) 
occurring at the Eocene-Oligocene boundary. In fact, when dealing with fauna 
having higher stratigraphic resolution (such as foraminifera), it has been 
demonstrated that extinctions and paleobiogeographic patterns are less 
instantaneous and more step-wise in nature than previously believed. 

Late Eocene Extinctions and Biogeographic Patterns 

Extinction of living plant and animal species is a process that has 
operated since life evolved on earth more than three billion years ago. It is a 
process that is of interest to scientists besides just paleontologists because the 
evolutionary process can provide important clues to the variation of 
environmental conditions throughout earth's history. This idea can be applied 
readily to paleoceanographic changes through time since the marine fauna and 
flora (and less directly, the terrestrial fauna and flora) all have certain 
environmental parameter tolerance ranges. If these environmental parameters 
vary beyond the adaptive capabilities of the organisms and species, extinction 


may occur. 


315 


Biologic extinctions may be categorized into two general types: 1.) mass 
extinction events, and 2.) normal or background extinction events. To be 
considered a mass extinction event, the extinctions should occur over a relatively 
short period of time (e.g., 1-5 million years), affect many different taxa, and the 
magnitude of the event should affect the taxa greatly (e.g., more than 50% of 
species go extinct). Without these characteristics, normal faunal turnover is 
considered background or normal extinction. 

As described earlier, the terminal Eocene extinction event is included in 
the marine fossil record at both the familial and generic taxonomic levels. The 
data presented by Raup and Sepkoski (1982, 1986) may not represent a true 
mass extinction event, however, when the timing of the event is considered. The 
data represent resolution only to stage level, with the mean stage duration given 
as 7.4 million years (Raup and Sepkoski, 1982). This means the timing aspect 
for the possible mass extinction may be too generalized when considering higher 
taxonomic levels, and interpretation of the cause and effect of paleoceanographic 
changes may be blurred when based on data such as these. Discussion of 
several taxa of marine invertebrates (in addition to the echinoids) and their 
extinction patterns near the end of the Eocene are described in the following 
paragraphs. 

Foraminifera have been studied in detail and used extensively for 
biostratigraphy of marine sediments. They are valuable (particularly the 
planktonic taxa) because of their wide distribution in ocean waters and their 
strong representation in many marine deposits. The resolution of foraminiferal 


316 


biozones based on various species can be defined within several hundred 
thousand years, so the timing aspect of their extinction patterns is excellent. 
Further, because this group contains many extant taxa, distribution patterns 
within the water mass and geographic distribution can be studied via living taxa 
to aid environmental interpretations for related fossil taxa. Specifically, 
geochemical analysis of their CaCOs tests yields isotope data that are valuable 
for these ecological changes in the water mass to be recorded and interpreted. 

Keller (1983) examined deep sea sediments from several DSDP sites in 
the Indian, southern Pacific, and southern Atlantic Oceans (sites 363, 292, 219, 
and 277). She made several interpretations of the water mass and climatic 
changes based on faunal turnover patterns of planktonic forams in the 
sediments. She noted four major changes in water mass stratification between 
the middle Eocene and late Oligocene based on global foraminifera faunal 
turnover. These changes in foraminifera species groups (where surface=warm, 
intermediate=cool, and deep=cold) suggest confirmation of the development of 
the psychrosphere (Benson, 1975) as the driving force for faunal turnover in the 
late Eocene. Furthermore, she noted that the global faunal turnover (extinctions) 
corresponds to major climatic change as well as a change in the global sea level 
curve. Flowever, it is stressed that the foraminifera faunal turnover at this time 
supports the model of catastrophic extinctions at the Eocene-Oligocene 
boundary, as previously proposed by other research. One important theme of 
Keller (1983) is the support of temperature change (cooling) as the driving force 


for the faunal turnover. 


317 


Keller (1986) also examined planktonic foraminifera in detail, with 
emphasis on the extinction events with regard to their timing, magnitude, and 
relationship to specific driving mechanisms. This study examined 37 late Eocene 
to Oligocene sections, most of which are DSDP sites from the Pacific, Indian, 
and Atlantic -Caribbean Oceans. Planktonic foraminifera datum events were 
established for dominant species and it was noted that 17 of 26 datum events fall 
between the Eocene-Oligocene boundary and 38.4 Ma, thereby reflecting rapid 
species turnover during a time of accelerated stepwise extinctions. 

Keller (1986) demonstrated that close examination of the foraminifera 
species record shows that successive extinctions occur abruptly over short 
stratigraphic intervals, creating a stepwise extinction effect. This is counter to the 
traditional view of a single, catastrophic mass extinction at the Eocene-Oligocene 
boundary. The number of species extinct at each stepwise extinction event 
generally is less than 15% of the species population and therefore would not 
represent a mass extinction. She points out, however, that the sum total of the 
late Eocene stepwise extinctions over the 3.4 million year interval (40.0-36.6 Ma) 
results in a nearly complete faunal turnover, with only 20% of the species 
surviving into the Oligocene. 

The Eocene-Oligocene extinction event in low to middle latitudes primarily 
represents a redistribution in the abundance of dominant species. Frequency 
changes in dominant species are less extensive at the boundary than at earlier 
stepwise extinction events. Temperature affinities of the dominant species 
indicate a gradual replacement of warm water species by cooler water species. 


318 


but Keller (1986) acknowledged that the extinctions may be the result of multiple 
causes. However, the proximal cause appears to be temperature-related 
(cooling) even if the ultimate cause is unclear. This idea is reinforced by other 
foraminiferal workers such as Berggren (1977), Corliss (1979), Keigwin (1980), 
Corliss et al. (1984), Keigwin and Corliss (1986), and Boersma et al. (1987), for 
both the gradual stepwise extinction pattern of foraminifera as well as the 
proposed causative force (temperature decrease). 

Deep-sea benthic ostracodes have been used to compare faunal changes 
with proposed paleoceanographic events for the Tertiary, and two events are of 
interest here. First, Benson (1975) used ostracode faunal shifts in deep-sea 
sediments to interpret the development of the psychrosphere at approximately 40 
Ma. Although limited extinction of ostracode taxa occurred, it was significantly 
earlier than the Eocene-Oligocene boundary event and Benson (1975) therefore 
called it the "40-million year event." Benson et al. (1984) examined in greater 
detail the global distribution and patterns of change for benthic ostracodes from 
156 DSDP sites (with specimens identified to the generic level). They found a 
decrease in global generic diversity and abundance in the ostracodes at the 
Eocene-Oligocene boundary (36 Ma), but it apparently was not as significant as 
the turnover at 40 Ma. Again, this shows an example of the potential error in 
assuming a mass extinction event at the end of the Eocene if the timing of the 
event cannot be carefully determined. Here, the ostracode event at 40 Ma likely 
would be included as part of the late Eocene extinction along with the true end- 
Eocene event (of lower magnitude) at 36 Ma. 


319 


Radiolarians also show faunal turnover at the end of the Eocene, but it is 
only of minor significance when compared to global diversity. Correlation of the 
radiolarian extinction pattern with impact horizons (microtektite layers) in sections 
from Barbados (Sanfilippo et al., 1985) and DSDP cores from the Gulf of Mexico 
and Caribbean (Glass and Zwart, 1977) have been documented. However, this 
faunal turnover may simply reflect part of a significant environmental change in 
the oceans as is interpreted from the foraminifera and ostracodes patterns 
(Keller, 1986). 

Molluscs also have been studied in some detail regarding extinction and 
relationship to causal factors such as shelf area and temperature changes, and 
impact events at the end of the Eocene (Hansen, 1987). Although Hansen’s 
study is restricted to Gulf Coast faunal patterns rather than global patterns, the 
pattern’s relationship to global climatic or paleoceanographic events may be 
interpreted from these data. Hansen (1987) compared molluscan extinction and 
diversification patterns from several formations of middle Eocene to Oligocene 
age. The taxa were identified to species level, but stratigraphic resolution was 
somewhat coarse since it was limited to appearance of taxa by formation. He 
found both gastropods and bivalves exhibited similar trends in diversity, with a 
high in the late middle Eocene, interpreted to likely be a function of increased 
temperature and shelf area at this time. 

The temperature drop present at the middle Eocene-late Eocene 
boundary and the Eocene-Oligocene boundary are strongly reflected in the 
molluscan diversity data. Hansen noted a molluscan species extinction of 86% 


320 


and diversity drop of 29% at the middle Eocene-late Eocene boundary and a 
97% extinction of the already depleted fauna at the Eocene-Oligocene boundary. 
He also compared species abundance of warm water genera versus cool water 
genera and he found the warm water genera suffer a higher specific extinction 
rate than cooler water genera across the Eocene-Oligocene boundary. This 
corresponds to similar results for planktonic foraminifera as reported by Keller 
(1983). A comparison of species-richness values for transgressive stratigraphic 
units versus regressive stratigraphic units shows no significant correlation 
between extinction of molluscs and shelf area for the middle Eocene through 
Oligocene units. Therefore, Hansen (1987) concluded that: 1.) the molluscan 
extinctions are stepwise in character through the late Eocene to early Oligocene 
(not catastrophic at the Eocene-Oligocene boundary) and, 2.) temperature drops 
appear to be the primary cause of extinction because of the selective loss of 
warm water taxa. 

Echinoid extinction and diversity patterns have been studied in 
relationship to temperature changes and sea level changes for the Tertiary, with 
particular emphasis on the terminal Eocene extinction event. McKinney and 
Oyen (1989) examined the statistical correlation of global and Coastal Plain 
echinoid diversity with two potential causative forces for extinction (i.e., sea level 
and temperature change) because these factors can be quantified from the fossil 
record. The resolution for the diversity data is limited to the stage level for 
Coastal Plain echinoid species and is even coarser for global species diversity 


321 


(epoch level), so this study resembles the resolution level used for the molluscs 
by Hansen (1987). 

A distinct pattern appears for the echinoid diversity at the Eocene- 
Oligocene boundary (for both global and Coastal Plain species diversity as 
reported by McKinney and Oyen, 1989), with a dramatic drop in diversity and 
increase in extinction. Comparison of the diversity data with sea level and 
temperature showed the affinity to temperature change to be stronger in a 
qualitative sense. This relationship was supported further after statistical 
correlation of these data reinforced the qualitative comparison quantitatively and 
they proposed that temperature may account for nearly 90% of the diversity 
variation. It must be noted that the data used by McKinney and Oyen (1989) are 
quite limited in stratigraphic and chronological resolution as compared with much 
of the foraminifera data, but the general interpretations of a significant 
environmental perturbance at the end of the Eocene agrees with other works 
described earlier. The limited resolution does not allow for further refinement of 
the timing of the paleoceanographic changes in detail, and therefore may or may 
not support the idea of a significant mass extinction event at the end of the 
Eocene. They also suggested that temperature may be only one of several 
forces causing extinction but it appeared to be a proximal cause even if not the 
ultimate cause of echinoid extinction. 

Data generated for this dissertation do not show a significant increase in 
the resolution of stratigraphic information or chronological information that would 
allow greater understanding of the Eocene-Oligocene boundary event. My data 


322 


tend to follow a similar diversity trend from the Eocene into the Oligocene 
(although slightly different in magnitude), with no significant diversity decrease 
from the middle Eocene into the late Eocene (as shown in some of the other 
taxonomic groups). Thus, at the resolution available for my data, it seems more 
probable that the environmental conditions were favorable throughout the late 
Eocene in this geographic region, but did indeed change significantly by the end 
of the Eocene. 


Chapter Summary 

The Cenozoic echinoderm record in Florida is relatively good, but certain 
aspects of collecting specimens (to aid in reconstructing the stratigraphic 
distribution and taxonomic diversity) can be greatly improved. One of the more 
valuable techniques which has allowed my research to increase the known 
diversity values for the Middle Eocene through the Pleistocene is close 
examination of the microscopic components of the sedimentary rocks sampled, 
both while in the field or after bulk-sampling and returning to the lab. These 
sieved samples have produced numerous echinoderm ossicles and other 
fragments that have proven to belong to taxa not recognized prior to my work 
and, in some cases, taxa that represent new species. A second method used in 
my research was the production of silicone peels of moldic rock units. Several 
taxa are known only from molds (either internal or external), and the peels made 
from them have reproduced the morphology in great enough detail to allow 
lower-level taxonomic identifications. 


As a result of closer examination of strata and their sedimentary rock 
during fieldwork, the study of a variety of museum and research echinoderm 
collections, and a thorough review of the published literature on Florida 
paleontology and stratigraphy, I have significantly improved what is known about 
the diversity and biostratigraphy of fossil echinoderms in the state. The resulting 
overall pattern of echinoid diversity from the Eocene through the Pleistocene is 
consistent with the previously described pattern throughout this time interval, 
except during the Miocene. My new stratigraphic and taxonomic records of 
echinoids show an increase in diversity for each epoch, but the pattern of change 
from the Paleogene to the Neogene now reflects a different trend. Previously 
published Florida echinoid data (McKinney and Oyen, 1989; McKinney et al., 
1992) show a diversity decrease from the Eocene to the Oligocene, and 
continuing into the Miocene. This contrasts with the worldwide echinoid diversity 
pattern, which shows a higher diversity in the Miocene than in the Oligocene. 

The revised Cenozoic echinoid diversity data from Florida show a trend that more 
closely follows (using proportional diversity of the raw number of species per 
epoch) the global pattern for echinoids. Flowever, when diversity values are 
normalized by epoch duration, the Florida data show a much lower proportional 
diversity for the Miocene as compared with the global pattern. 

Florida is not the only place with this low Miocene normalized diversity, as 
a similar pattern also is present in Jamaican echinoids (Donovan, in press), 
which may represent a regional effect in the Caribbean and Gulf of Mexico. One 
of the reasons for the improved alignment of local (i.e., Florida) and global 


323 


324 


patterns is the significant number of additional taxa recognized from the Miocene 
as a result of this study. The Cenozoic echinoids in Florida still do not directly 
mimic the global pattern, but as work continues the local, regional, and global 
patterns may begin to match more closely. 

The Florida echinoderm diversity pattern and fossil record is influenced by 
a number of biases, not unlike the fossil record of other taxa. These biases 
include factors such as variations in outcrop exposure among the Cenozoic 
formations, revisions of stratigraphic boundaries and names, differential 
preservation potential in carbonate versus siliciclastic environments, unequal 
temporal ranges of epochs, revisions of taxonomy at the generic and specific 
levels, and the normal tendency for people to collect unbroken and larger fossils 
preferentially over small, fragmented, or disarticulated skeletal remains. Any of 
these biases, whether independent or additive, affects the ability of 
paleontologists to assemble a complete picture of the echinoderm record from 
Florida or any other geographic region in the world. This dissertation is only the 
first attempt to synthesize a complete Cenozoic echinoderm database for Florida, 
and represents a building block which I use to augment the fossil echinoderm 
record in the state and move forward in the research on the echinoderms. 


CHAPTER 5 

ALLOMETRIC HETEROCHRONY IN MELLITID ECHINOIDS: 
A CASE STUDY FROM FLORIDA 


Preface to the Biometric Analysis 
Original Research Objectives 

The original objective of my research was to carefully study the growth of 
modern echinoids with the goal of determining if growth in echinoids is such that 
skeletal calcite accurately records the individual’s ontogenetic age. This would 
permit standardization by common age in evolutionary series of fossil lineages so 
that heterochronic patterns and mechanisms could be assessed. However, the 
growth analysis was unsuccessful largely because of mass mortality of the 
modern test population of echinoids before a full year of growth was completed. 
Another unexpected result was the lack of visible and traceable growth lines in 
the skeletal plates of those individuals that survived for the necessary full-year 
time period. Thus, quantitative analysis of the biometric data gathered from 
selected echinoids of Florida proved to be the most problematic for me to 
complete in the method anticipated in the planning stages of the dissertation. 

Growth in echinoids occurs by peripheral accretion of individual test plates 
as well as addition of new plates (Swan, 1 966; Raup, 1 968; Smith, 1 984). 


325 


326 


Unfortunately, little is known concerning the growth rate of echinoids during 
ontogeny. Groups such as molluscs (Jones, 1988) and plants (Guerrant, 1988) 
allow determination of their ontogenetic age with fine resolution. Thus, we may 
assume reasonable confidence for interpretation of the mechanism responsible 
for morphological change in the taxa. For many other taxa (including the 
echinoids) a technique for determining an organism's ontogenetic age is yet 
unproven. 

Growth Study Procedure 

Determination of growth line regularity was attempted by “tagging" living 
echinoids via injection with tetracycline hydrochloride. The echinoid species 
Mellita quinquiesperforata (irregular echinoid) is found living along the Gulf of 
Mexico coast of Florida in the Cedar Key and Seahorse Key area. Following the 
tetracycline tagging technique of Kobayashi and Taki (1969), a large number of 
individuals of the species were collected and tagged. I followed the optimal 
dosage of the tetracycline (determined by Kobayashi and Taki) as 1-2 mg per 10 
g live urchin weight, with the tetracycline mixed with seawater in a ratio of 1 mg 
per 0.1 ml of seawater, and the solution was injected into the urchin through the 
peristome. 

After the injection of tetracycline, the echinoids were placed in "fenced" 
enclosure areas near Seahorse Key that allowed continued exposure to natural 
conditions, yet also allowed the tagged individuals to be collected later (while 
minimizing specimen dispersal). The enclosures consisted of hardware cloth 


327 

(~12 mm mesh “chicken wire”) fence material embedded 10-20 cm into the 
substrate to prevent escape of the sand dollars. 

The tetracycline is incorporated into the calcite test as it accretes (at least 
ideally this is true) and results in distinctly colored lines in the test when the test 
plates are viewed under reflected ultraviolet light. Therefore, the injection serves 
as a time reference marker in the test's plates and potentially can be used to 
interpret periodicity of growth line development. A number of the previously 
tagged individuals then were collected at three month intervals, measured, and 
sacrificed to determine any seasonal growth variation and to document the 
presence of any new growth lines. Also, environmental conditions such as 
salinity and temperature were recorded when collecting the specimens. 

Two problems occurred during this experimental procedure that required 
me to alter the dominant focus of my research from that of heterochrony analysis 
to biostratigraphy and diversity pattern analysis. The first problem encountered 
was the mass mortality of the study population of echinoids. As part of the 
procedure for growth study of the population, I traveled to Seahorse Key at 
intervals of approximately three to four weeks to gather data of general test 
length and width biometrics, as well as measurements of salinity and water 
temperature conditions. At approximately eight months into the 12-month growth 
study I discovered that nearly all echinoids in the enclosures had died. A 
significant amount of algae as well as other organic and non-organic matter had 
become entangled in the echinoid enclosure fencing material through some 
event such as storm wave activity. These attached materials thereby acted as 


328 


baffles preventing water circulation through the enclosures, and initiating the 
settlement of a thick (4-8 cm) layer of clay- and silt-sized sediment out of 
suspension. Therefore, I interpret the mortality of the echinoids to be the result 
of “death by suffocation,” since these organisms generally require coarser 
substrates with modest water movement to allow efficient respiration of dissolved 
oxygen via their tube feet. 

The second problem involved analyzing growth lines in the echinoid test 
plates. Since the mass mortality occurred before a full year had passed, the 
ability to judge periodicity of growth line emplacement became undeterminable. 
Growth lines in echinoid test plates were reported as early as the mid-1800s in 
the literature, and the possibility these lines are cyclic in formation is both 
supported (Raup, 1968; Jensen, 1969; Weber, 1969; Pearse and Pearse, 1975; 
Smith, 1984) and disputed (Heatfield, 1971; Ebert, 1986). Supporters of cyclic 
growth line development generally attributed their formation to seasonal cycles in 
growth rate due to factors such as temperature, reproductive state, food 
availability, or storms. Studies such as Ebert (1986) and Weber (1969) were 
concerned with growth lines in spines rather than test plates. Ebert believed 
these rings were due to disturbance effects as well as growth and thus are not 
cyclic in nature. Weber found rings established even when trauma was not a 
factor. Neither worker was able to quantify the period of time present between 
ring establishment. Therefore, it was impossible to distinguish the chronological 
significance of any lines in the plates. Furthermore, when the plates of the 
“early" mortality echinoids were examined for tetracycline banding, no 


329 


consistency in the age marker bands could be established. That is, I could see 
fluorescence of the tetracycline under ultraviolet light, but it was distributed 
sporadically rather than consistently through various plate sections in the test. 

The effort was not a complete failure, however, as I have incorporated the 
biometrics of the Seahorse Key echinoids into the database of biometrics from 
the fossil specimens. Mellita quinquiesperforata is reported herein as part of the 
fossil record of Florida, but the recovered fossils are poorly preserved (usually 
highly fragmented) and therefore did not allow measurements to be made on 
those specimens. As a result, I was able to use the biometric data from these 
specimens as part of the allometric heterochrony analysis of the mellitids in 
Florida. 


Introduction and Heterochrony Overview 

Heterochrony, the change in timing of an organism's development during 
ontogeny, is not a new idea in the field of evolutionary study. Ernst Haeckel 
(1875) provided the term to delineate anomalies in evolution as described in his 
biogenetic law regarding single organisms (within a species). Our present 
concept of Haeckel's idea was indelibly altered by de Beer (1930), who 
generalized it to become the change in developmental timing of a feature relative 
to the same feature in an ancestor (see Gould, 1988). 

Perhaps it is true that the specific definition of heterochrony accepted and 
applied today is not what was intended by the original presentation of Haeckel. 
Nonetheless, heterochrony can give useful insight to ecological changes that 


330 

influence these developmental changes. As McNamara (1982) noted, these 
changes may result in normal phenotypic variation or, if of adaptive significance, 
selection for the morphologic variant leading to speciation. 

An increasing number of publications regarding heterochrony in the 
evolution of various taxonomic groups reflects the increasing awareness and 
interest in this field in recent years. Two publications in particular, Gould (1977) 
and Alberch et al. (1979), have elucidated the important tripartite relationship of 
time (age), size, and shape to heterochrony. Gould's approach is more 
qualitative while Alberch et al. emphasize a quantitative analysis of the 
heterochronic processes and their identification. McNamara (1986) provided a 
concise yet valuable explanation of the terminology and diagnostic 
characteristics for the heterochronic processes found in current literature (Figure 
5-1). More recently, McKinney and McNamara (1991) have elaborated on the 
mechanisms, modes, and effects of the evolutionary changes resulting from 
heterochrony as related to wide variety of living organisms. 

Allometry has been stressed in previous heterochronic studies, typically 
through bivariate morphometric analysis of ontogenetic growth trajectories. 
Ontogenetic age data for fossils often is not obtained or is disregarded and 
heterochrony is then described using only two of the three characters, size and 
shape change. It is not possible to characterize the change in developmental 
timing unless the ontogenetic age is known at the various developmental stages 
for the individual. When size is equated with age, "allometric" heterochrony 
rather than true heterochrony is defined (McKinney, 1988; p. 24). That is. 


HETEROCHRONY 


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allometric heterochrony is a pattern interpreted via analyses involving shape and 
size, rather than ontogenetic age. Organisms that have continuous grov\/th 
throughout ontogeny should reasonably be expected to reflect age via body size. 
It must be noted that problems arise, ho\A/ever, in cases where body size is not a 
reliable indicator of ontogenetic age, and thereby invalidate the size-based 
approach to heterochronic analysis (see McKinney, 1988; McKinney and 
McNamara, 1991 for further explanations). An example of an invertebrate group 
whose body size does not necessarily reflect a direct correlation with age are 
bivalves, which may have two individuals of the same size varying in ontogenetic 
age by 50-100 years (Jones, 1988). Recently, Jones and Gould (1999) 
expanded upon the implications and explained how the assumption of size as an 
indicator of age has produced errors in characterizations of heterochrony 
patterns for Grvphaea . Thus, I recognize the limitations inherent in size-based 
interpretations and accordingly consider my results to be "allometric" 
heterochrony interpretations in contrast to those based on known age (i.e., the 
"true" heterochrony styles). 

The purpose of the analysis described herein is to illustrate the styles of 
allometric heterochrony present in select species of sand dollars in the family 
Mellitidae from the Cenozoic in the southeastern U.S.A. The data included within 
this chapter are the only data for this dissertation that include fossils not 
necessarily present in strata from the state of Florida (those few selected fossils 
were originally collected from South Carolina localities). The data generated 
from the species of Encope and Mellita are derived primarily from fossils. Mellita 


333 


guinquiesperforata is the only species represented by individuals sampled from a 
modern population (rather than a fossil population) located surrounding Seahorse 
Key, Florida, in the Gulf of Mexico. Bivariate regression analysis was completed 
on morphologic characters to generate allometric grov\/th trajectories used for the 
heterochrony analysis of these mellitids. Finally, a brief comparison v\/ith 
paleoecological and modern ecological conditions during the species' evolution is 
provided to help identify the potential environmental selective impetus in the 
evolutionary history of the fossil mellitids. 

Materials and Methods 

Materials Examined 

The primary goal of this project is to identify the styles of evolution present 
in available species of fossil mellitids from the southeastern Coastal Plain, and in 
particular, Cenozoic fossils from Florida. The species under examination are 
those that are reasonably v\/ell preserved, thereby allowing easy specimen 
preparation and access to the variety of morphological characteristics for 
biometric purposes. Second, the species were available for study either via 
personal collection through fieldwork or through examination in museum 
collections. Up to nine species (depending on taxonomic validity of species 
designations) of mellitids are included in my database of biometric 
measurements and analyses. 

The species examined here include the fossils Encope tamiamiensis 
Mansfield, 1932 (Tamiami Formation, Pliocene, Florida), Encope michelini 


334 


Agassiz, 1841 (uncertain unit ?, Pleistocene ?, South Carolina; also found in the 
Anastasia Formation, Pleistocene, Florida), and Mellita aclinensis Kier, 1963 
(Tamiami Formation, Pliocene, Florida). The fourth taxon is represented by 
individuals of Mellita quinquiesperforata (Leske, 1778) that were sampled from a 
modern population near Seahorse Key, Florida, in the Gulf of Mexico (note that 
this species also occurs as a fossil in the Satilla Formation of Florida). Personal 
fieldwork provided all specimens of M- quinquiesperforata while all specimens of 
M. aclinensis and E. tamiamiensis are part of the Invertebrate Paleontology 
Collection at the Florida Museum of Natural History in Gainesville, Florida. 

Finally, all measured samples of E. michelini were obtained from the U.S. 
National Museum in Washington, D.C. 

Data Acquisition Methods 

Biometric data were gathered using Fowler Ultra-Cal II electronic calipers, 
accurate to 0.01 mm. The calipers were connected to a Dell NL20 laptop 
computer, which allowed the each measurement's datum to be sent and 
compiled directly in a spreadsheet program running on the computer. Following 
collection, data were then exported to statistical analysis programs to complete 
the data analysis. Univariate statistics were calculated using a Macintosh 
computer and the commercial statistics computer program StatView (both 
versions II and 5.01 were used). All biometric measurements were subjected to 
base 10 logarithmic transformation before regression analysis. Calculation of a 
Z-statistic, as a test for determining significant differences in slope means 


335 


between species pairs, and conversion of least-squares regression output to 
reduced major axis regression (RMA), were completed using a PC computer and 
computer programs I wrote via the BASIC programming language to produce 
such conversions. 

Biometric measurements were produced using 372 individuals from 
several echinoid species within the family Mellitidae. The species that were more 
abundant in the collections allowed more than 100 specimens to be measured. 
Several species are relatively rare in the field as well as in fossil collections, and 
therefore are represented by as few as 7 specimens in this data set. The 
number of specimens measured for each species in this study ranges from 7- 
131. Up to 64 separate measurements were taken on each echinoid specimen, 
depending on the state of preservation of the individual fossil. All trait 
measurements were taken on all specimens, except in those circumstances 
where one or more species did not have the same morphologic feature present 
as part of its skeleton. For example, ambulacral lunules are not present in 
species of Encope , therefore the trait has no data available for those specimens. 
Incomplete specimens, or those fossils with encrusted or cemented matrix that 
was unable to be removed, had fewer biometric data compiled from them. 
Morphologic landmarks were established to allow similar data files to be 
established for each species, and include common measurements such as test 
length (TL), test width (TW), peristome length, width, and position (PSL, PSW, 
PSP), among numerous other traits discussed below. 


336 


Biometric Traits Evaluated 

Biometric data were gathered from morphologic traits present on all 
species of mellitids examined in this study. Many of these traits commonly are 
used in biometric analyses of modern and fossil echinoids, while several traits 
and associated variables were established as unique to this project. Morphologic 
landmarks were defined to allow similar data files to be established for each 
species. Trait types and locations include most of those described by Harold and 
Telford (1990), as well as additional variables chosen as unique for this study. 
Figure 5-2 shows representative locations of biometric measurements associated 
with the species of mellitids while Table 5-1 lists the trait abbreviation or acronym 
and the description of the trait location on the echinoid test. 

All numbering associated with traits (using l-V and 1-5) follows Loven's 
(1892) system, in which ambulacra use Roman numerals and interambulacra use 
Arabic numerals. Most traits identified and measured are in multiples of five due 
to their association with the ambulacra and interambulacra on the echinoids. 
Therefore, those trait measurements are designated by having the affiliated 
Arabic and Roman numerals (in parentheses after the trait acronym) using the 
Loven system as noted above. Biometric traits that are not associated with the 
ambulacra and interambulacra skeletal regions lack such numerals. A total of 24 
independent and unique morphologic traits were evaluated, that resulted in a 
sum total of 64 traits when the five-fold multiples are considered as individual 


traits. 


337 


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338 


Table 5-1. 

Biometric trait descriptions. 

TRAIT 

TRAIT DESCRIPTION 

TL 

TW 

PSL 

PSW 

PSP 

Test length 

Test width (at midpoint of TL) 

Peristome length 
Peristome width 

Peristome position (from anterior test margin to anterior peristome 
margin) 

PPL 

PPW 

PPP 

Periproct length 

Periproct width (at midpoint of PPL) 

Periproct position (from anterior periproct margin to anterior 
peristome margin) 

POSAP 

Position of apical system (from anterior test margin to center of 
apical system) 

ANLL 

ANLW 

Anal lunule length (interior lunule length, aboral side) 

Anal lunule width (interior lunule width, aboral side, at midpoint of 
lunule length) 

ANLP 

Anal lunule position (distance from anterior lunule margin to apical 
system center) 

PD(I-V) 

TWMX 

LTH(1-5) 

Pressure drainage channel span (max. width, adoral surface) 

Test width maximum 

Longitudinal test height (at 5 equidistant points starting at anterior 
test margin) 

TTH(1-5) 

Transverse test height (at 5 equidistant points starting at left test 
margin) 

PAL(I-V) 

Petaloid ambulacrum length (aboral ambulacrum length, from apical 
system margin to maximum pore-pair position) 

PAW(I-V) 

Petaloid ambulacrum width (aboral ambulacrum width, at midpoint 
of ambulacrum length, from outer pore-pair to outer pore-pair) 

THMX 

AL(I-V) 

Test height maximum 

Ambulacrum length (adoral surface, peristome margin to test 
margin) 

IL(1-5) 

Interambulacrum length (adoral surface, peristome margin to test 
margin) 

ALL(I-V) 

ALW(I-V) 

Ambulacral lunule length (aboral surface, interior length) 

Ambulacral lunule width (aboral surface, interior width at midpoint of 
length)) 

ALP(I-V) 

Ambulacrum lunule position (from center of apical system to 
adapical lunule margin) 


339 


Data Analysis Methods 

Summary statistics were generated for all biometric traits to use as 
reference for comparison with original species descriptions and morphology 
observed in populations that were sampled. Bivariate regression analysis was 
applied to the data to generate growth trajectories for the biological characters 
used as variables. Reduced major axis (RMA) regressions were produced and 
used here rather than least-squares regressions because the RMA method is 
more appropriately applied to biological growth systems where a truly 
independent variable may be difficult to discern (see Davis, 1986 for a review of 
details regarding variable assumptions in this method). Since all components 
(i.e., morphologic traits) of this analysis involve allometric changes during growth 
of the echinoids, a true, independent variable may not exist and therefore the 
RMA method is a better regression option. Recognizing this factor, test length 
(TL) was chosen as the most likely representative or proxy of body size, and 
therefore is designated as the independent (x-axis) variable in all calculations. 
This regression method reveals allometric changes that occur as part of the 
skeletal growth of the echinoid species, which in turn then is used to interpret 
allometric heterochrony styles present in the taxa of interest to this study. 

McKinney (1986) established models for allometric heterochrony 
interpretations based on regression data for biometrics. The six styles of 
heterochrony may be distinguished based on the slopes and y-intercepts for the 
equation of generated regression lines, as well as species body size. An 
illustration of this model's format is provided in Figure 5-3. Explanations of all 


340 


assumptions and limitations in using this diagnostic model are provided in 
McKinney (1986, 1988) and McKinney and McNamara (1991). Regression data 
from the selected Neogene mellitids then vjere compared with the model for 
interpreting heterochrony patterns present in the taxa. Approximately 3500 
bivariate regression plots were generated during my analysis and examples of 
two such plots are provided in Figure 5-4. It is possible to visually distinguish the 
heterochrony pattern from plots such as these, but statistical tests were used to 
confirm any interpretations. Due to the large amount of statistical output (both 
graphic and tabular) generated during the analysis, I only present a simple 
example of the output here and summarize the rest of the information in tables in 
the results section. 

In this paper, RMA slopes and y-intercepts are compared at the .05 level 
of significance using the Z-test statistic, and all regressions must have r^=0.80 or 
greater for inclusion in the comparisons. Regressions for all data included here 
use log-transformed biometric data. Finally, where interpretations of progenesis 
or hypermorphosis are provided, the body size (TL) critical value is determined 
using the TL value at two standard deviations above the mean for the species 
sample (Table 5-2). 


Results 

The general results of the heterochronic relationships between the species 
pairs are shown in Table 5-3. The method used to determine the evolution style 
follows that used by McKinney (1988). A clear dominance of peramorphosis 


341 


PROGENESIS HYPERMORPHOSIS 




PREDISPLACEMENT POSTDISPLACEMENT 




ACCELERATION 



S 


NEOTENY 



S 


Figure 5-3. “Allometric” heterochrony, as classified by the ontogenetic plots of 
related species. Axis S = body size (test length for the echinoids), 
and axis T = trait measurement. S and T do not require variables to 
be log-transformed for the model to work, but the data in this 
dissertation are log-transformed for the analysis. Sketches illustrate 
hypothetical body size (light circle), and trait size (e.g., test width, 
ambulacrum length, etc.). (From McKinney, 1988) 


Figure 5-4. Examples of reduced major axis (RMA) regression plots for the 

species pair M. aclinensis and M- auinguiesperforata . Data is log- 
transformed and the independent variable (proxy for size) is the test 
length (TL). 

A) Dependent variable is anal lunule position (ANLP) and, based on a difference 
in RMA slopes, the heterochrony style illustrated is acceleration (slope of 
descendant M. auinguiesperforata is less than ancestor; Z=-8.58, n=1 1 9 for 
M. aclinensis and n=7 1 for M- auinguiesperforata ) . 

B) Dependent variable is test width (TW), and since both the slope and y- 
intercept are not statistically different but the mean TL value is greater for the 
descendant, the heterochrony style illustrated is hypermorphosis (Zsbpe=-0.31, 
Zy-intercept=0.90, n=117 for M. aclinensis and n=64forM- auinguiesperforata) . 


343 



log(x) of TL 

y = 0.74x + 0.98, = 0.921 M. aclinensis 

y = 1.25x 0.40, = 0.821 M. Quinauiesperforata 



y * 0.99x + 0.006, = 0.997 aclinensis 

y = I.OOx - 0.014, 1 ^ = 0.978 M. quinquiesperforata 


Table 5-2. Echinoid body size (TL) calculations for mellitid species included in heterochrony analysis The following represents the 
summary data for the echinoids regarding length, maximum length, and length at 2 standard deviations above the 
mean test lerigth (both for corrected, where n<10, and uncorrected factors, where n 10). Asterisks indicate a modified 
file that includes closely related species that may be conspecific, rather than truly unique. Acronyms included in 
parenthesis beneath species names are used as code for location within biometrics data file of Appendix. 


344 


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E. michelini 9 81.83 12.62 11.32 97.02 104.48 


345 


exists in the ancestor - descendant pairs, with paedomorphosis only accounting 
for less than 20% of the traits involved in the analyses. An unequal balance 
exists among the three styles of peramorphosis as well, with acceleration 
distinctly more common in the M. aclinensis -M. quinquiesperforata and E. 
tamiamiensis -M. quinquiesperforata pairs. Hypermorphosis is most common 
among the morphologic traits of the E. tamiamiensis -E. michelini relationship, 
and it is secondary in prevalence to acceleration in the other species pairs 
illustrated here. The other style of peramorphosis, pre-displacement, is relatively 
rare or absent for each of the three species pairs. A point that may be made 
regarding the heterochrony styles is the dissociated rather than global nature 
within the individuals of a given species. This is not unique to the mellitids, or 
echinoids as a group, and should be expected when a fixed number of 
morphologic traits are varying during allometric growth of the organism (i.e., a 
"closed" system of sorts). 

An illustration of how the most common varieties of peramorphosis can be 
readily determined graphically is shown in Figure 5-4 for two of the morphologic 
traits analyzed in the M. aclinensis - M. quinquiesperforata relationship. This was 
done for all morphologic traits examined in each species pair (though not shown 
for all traits), and then related to the graphic model of McKinney (1986). It is a 
relatively simple procedure and model to use and visually perceive when the 
slopes of various regression lines are distinctly different, as well as in the cases 
where the slopes and y-intercepts of regression lines are not different, as shown 
in the two cases in Figure 5-4. All equations of the regression lines were tested 


346 


for statistical significance (at the 0.05 confidence level) in difference in slope and 
y-intercept for all traits (using the Z-test), no matter how easy or difficult to 
perceive by visual methods alone. It must be noted that for this study, no 
cladistic phylogeny was completed, and therefore the ancestor-descendant 
species pairs must be viewed with caution until such data are available. 

The most interesting aspect of these heterochrony patterns is derived not 
from the simple documentation of the patterns, but rather by relating the patterns 
to potential "causal" forces or environments that allow selection for traits to occur. 
McKinney (1986) demonstrated a non-random relationship between the energy 
of environment in which certain fossil echinoid taxa were living and the 
heterochrony patterns of evolution among closely related taxa. For the echinoids 
he examined (none of which were mellitids), he found a high proportion of 
species-pairs reflecting the presence of paedomorphosis (particularly neoteny) 
when evolution occurred in a change from unstable to more stable ecologic 
conditions. The opposite held true in which patterns of peramorphosis reflected 
a change from stable to more unstable environments. 

The evolutionary trends of the mellitids examined in this study seem to 
support the model McKinney proposed. When the evolution of the species pair 
M. aclinensis - M. quinquiesperforata is considered, I have shown a 
predominance of peramorphosis styles of acceleration and hypermorphosis for 
these species. The fossils of M- aclinensis are found in great abundance in 
portions of the Tamiami Formation (Pliocene) at some of the classic Pliocene 
shell pits of central and southern Florida. The paleoecology of the beds 


347 


containing these echinoids at a locality such as the Lomax-King Pit (now inactive) 
in Charlotte County can be interpreted as representing intermediate to shallow 
water depths and with moderate water energy. The modern environment where 
the extant species of M. quinquiesperforata was collected, represents an 
intertidal to very shallow subtidal water depth, having a moderate- to high-water 
energy (particularly during storm events). The fossil representatives of M. 
quinquiesperforata also have been collected from high-energy nearshore sand 
deposits of the Satilla Formation. Although the two species do not represent 
extreme differences in water energy and depth, based on my paleoecologic 
interpretation it is likely that modern environment is a higher energy setting on 
average than that which existed at the time M. aclinensis was inhabiting the 
paleo-coastline of Florida. 

Although not all species measured are represented in the examples of this 
chapter, the general pattern is present throughout the species of the family 
Mellitidae. As my summary of the heterochrony analysis, I interpret that the 
mellitid echinoids examined as part of this study tend to follow the general 
echinoid model and pattern of evolution shown by McKinney (1986) and 
McNamara (1982). The mellitids produce a general peramorphocline trend in the 
southeastern Coastal Plain, shifting from slightly deeper and more moderate 
energy water conditions, to shallower depths and higher energy. During this 
evolutionary transition, the mellitids exhibit predominant patterns of 
peramorphosis, with acceleration and hypermorphosis styles most common. 


348 


Table 5-3. Percent distribution of allometric heterochrony styles for selected 
ancestor-descendent species pairs as interpreted from RMA 
regression analysis. Note predominance of peramorphic styles for 
each species pair. Regression lines must be at r ^ = 0.80 or greater 
for inclusion in final analysis. Thus, less than 60% of total traits 
measured are included in these calculated values. 


PAEDOMORPHOSIS PERAMORPHOSIS 

Mellita aclinensis vs Mellita quinquiesperforata 
Neoteny Acceleration 


1 trait 2.7% of total 

1 8 traits 48.6 % of total 

Progenesis 

0 traits 0.0% of total 

Hypermorphosis 
1 1 traits 29.7 % of total 

Post-Displacement 
6 traits 16.2 % of total 

Pre-Displacement 
1 trait 2.7% of total 

18.9 % of total traits 

81.0 % of total traits 


Encope tamiamiensis vs Encope michelini 


Neoteny 

Acceleration 

3 traits 9.7% of total 

10 traits 32.3 % of total 

Progenesis 

0 traits 0.0% of total 

Hypermorphosis 

18 traits 58.1% of total 

Post-Displacement 
0 traits 0.0% of total 

Pre-Displacement 
0 traits 0.0% of total 

9.7 % of total traits 

90.4 % of total traits 


Encope tamiamiensis vs Mellita quinquiesperforata 


Neoteny 

Acceleration 

3 traits 7.9% of total 

21 traits 52.6 % of total 

Progenesis 

0 traits 0.0 % of total 

Hypermorphosis 
8 traits 21 .0 % of total 

Post-Displacement 
4 traits 10.5% of total 

Pre-Displacement 
3 traits 7.9% of total 

18.4 % of total traits 

81.5 % of total traits 


CHAPTER 6 

SUMMARY AND CONCLUSIONS 


The Cenozoic fossil record of Florida is renowned world-wide for the 
abundance of material, the diversity of taxa, and the wealth of paleontological 
and paleoecological information that can be collected from the strata. The fossil 
invertebrate record is particularly good, especially for taxonomic groups like the 
molluscs and foraminifera, with some of the Neogene beds composed almost 
entirely of shell material. Fossil echinoderms are a significant component of 
several formations, including the Ocala Limestone and Suwannee Limestone 
from the Paleogene and the Tamiami Formation from the Neogene, while other 
units apparently had only sparse distributions according to the published record 
through 1994. My research has significantly changed our understanding of the 
diversity pattern of echinoderms, particularly in the transition from the Paleogene 
to the Neogene. 


New Echinoderm Diversity Patterns 

Most of the echinoderms reported as fossils are echinoids, with only very 
limited reports and descriptions of asteroids, ophiuroids, and crinoids. Several 


349 


350 


new reports of ophiuroids and asteroids are included herein (see chapter 3), 
though most identifications are very limited since isolated ossicles are not 
normally reliable guides for lower taxonomic identifications. Most of the changes 
in diversity involve the echinoids as a group. When this project was started, the 
total echinoid diversity from the Middle Eocene through the Pleistocene was 68 
species, whereas I now report the diversity to be 100 species. This represents 
an increase of nearly 50% over the previously published diversity, with the new 
records and new reported occurrences now accounting for 32% of the total 
record. The additional taxa reported and counted in this analysis are both new 
stratigraphic records for the state as well as any fossils I interpret to be new 
taxonomic species. Therefore, not all of the new additions are new taxa in an 
evolutionary context, yet even the newly reported occurrences of species are 
important to more fully understand the patterns of change temporally as well as 
spatially. 

Another important change in echinoid diversity as a result of the research 
completed herein occurs across the Paleogene-Neogene boundary. The 
previously published diversity values were at eight species in the Oligocene and 
five species in the Miocene. I generate a much different pattern based on my 
revised data, so that the diversity increases from 1 1 species in the Oligocene to 
22 species in the Miocene. This reflects a dramatic increase in diversity that I 
propose to be the primary result of collecting style rather than principally 
controlled by extinction, origination, or an environmental cause. Most of the taxa 
considered herein as new stratigraphic records and new taxonomic records in the 


351 


Miocene likely were recorded simply due to sampling of fragmented, incomplete 
specimens. Most paleontologists are interested in more complete specimens for 
biometric analysis or descriptive work. Fragmented fossils do not allow easy 
identifications to be completed and thus may not lend themselves as well for 
application to community analysis, paleoecology reconstruction, or species 
descriptions. Unfortunately, this has resulted in reported diversity of the Miocene 
to be very low, but clearly this is not true when small size-fractions or fragmented 
specimens are examined and documented for the fossil record. 

I believe several other biases (in addition to collector bias) have 
contributed to the previously reported low diversity of the Miocene as well as the 
overall diversity pattern in the Cenozoic echinoids of Florida. The status of 
stratigraphic nomenclature and how such strata are defined greatly affects the 
values attributed to “new stratigraphic records” in the state. If one formation is 
subdivided into two distinct units, the echinoderms present in at least one of 
those units would, by definition, become a first reported occurrence of the 
species within that formation. Conversely, if consolidation of stratigraphic units 
occurs, it is possible that a species may be removed from the biostratigraphic 
distinction of a new report for a formation. 

Mineralogical composition of the rock units can also influence the diversity 
pattern due to its influence on weathering and preservation potential of the 
fossils. In Florida, the general trend is for better preservation of echinoderms in 
carbonate rock units of the Paleogene and poorer preservation styles in Neogene 
siliciclastic units. The carbonate rock layers buffer acidic groundwater solutions. 


352 


thereby reducing dissolution or test degradation when the fossils is surrounded 
by the limestone sediment. Siliciclastic units, on the other hand, do not have an 
abundance of minerals that can buffer acidic solutions. This results in more 
effective chemical degradation by the acid when a carbonate fossil grain, such as 
tests and ossicles of echinoderms, comes in contact with acidic groundwater. 

The chemical reaction need not be complete in order to influence the physical 
strength and durability of the skeletal grain; partial dissolution may be enough to 
enhance the extent of breakage. This may help explain the predominance of 
fragmented and highly weathered fossils of all varieties in selected siliciclastic 
beds of Neogene units. 

Several other biases affect the diversity record of echinoderms in Florida 
(discussed in Chapter 4), but perhaps not to the extent that sampling bias, 
stratigraphic nomenclature changes, and the diagenetic setting may have on the 
record. Regardless, one must be aware of such biases to accurately interpret 
the patterns present in the fossil record. 

Taxonomic ImpJications 

Another important result of the work completed is the collection of 
numerous fossils that appear to be new species of echinoderms. Relatively few 
specimens from Paleogene formations may be undescribed taxa, but the 
Neogene units (particularly of Miocene age) may have more than eight echinoid 
species awaiting formal description. In addition to the echinoids, several asteroid 


353 


and crinoid specimens remain unidentified to specific level and are possible new 
taxa. 

The higher proportion of potential new species in the Paleogene likely is a 
function of few paleontologists closely examining the broken and disarticulated 
material associated with the formations. Descriptions of incomplete specimens 
often result in rather tentative identifications, so it is possible that even if such 
taxa had been collected in the past, they were discarded rather than described. 

In more recent years, technology improvements have provided the necessary 
tools to examine fragments in much greater detail. Scanning electron 
microscopy allows observation of ossicle and skeletal plate micromophology that 
are often distinctive for identification to lower taxonomic levels. Therefore, even 
the apparently “useless” isolated plate or radiole fragments may give clues to 
family, genus or species identification (see Donovan and Carby, 1989; Gordon 
and Donovan, 1992; and Dixon and Donovan, 1998 as examples). 

Finally, biometric analysis of the selected mellitid species indicated 
several species may be too similar to justify unique species status. As an 
example, Encope macrophora (a species found in North and South Carolina) and 
Encope tamiamiensis (a species found in Florida) have nearly identical 
regression line slopes when various physical traits such test width are regressed 
with test length. The growth trajectories for such traits often are not statistically 
significant, and even when a statistical difference exists, one cannot discern a 
difference when looking at the hand sample. This tendency to name new 
species based purely upon geographic differences among populations rather 


354 


than significant morphological variance creates turmoil in the realm of taxonomic 
worker. I believe significant work can be done in the future to help clarify and 
resolve problems like this by combing statistical analyses with detailed qualitative 
characterizations. 


What Work Lies Ahead? 

I believe my work in biostratigraphy has just begun, and one of the most 
important components of the echinoderm biostratigraphy is to refine the 
stratigraphic resolution of the data I have gathered thus far. Most of the data are 
limited to formation or, in far less common cases, members of formations. 

Isolated data exist with measured section information available, but it is not 
common enough for most detailed tasks. My goal is to continue sampling 
stratigraphic units, looking for any fragmented or complete fossil material that 
may produce indications of greater diversity in the Florida units than we currently 
recognize. Measured sections, with lithologic descriptions and (ideally) 
petrography will allow enhance facies and biofacies distributions to be interpreted 
and utilized for paleoecology purposes. 

A second component that I believe needs to be investigated is a broader 
picture of the echinoderm biostratigraphy. Cooke’s work (1959) describing the 
Cenozoic echinoderms of the southeastern U.S. is over 40 years old. Much of 
his taxonomic work is out of date with respect to stratigraphic and geographic 
distributions of the echinoids. Numerous taxonomic revisions to his “cook(e)book 
of echinoid paleontology” require the database to be updated, and no one thus 


355 


far has compiled enough new data to make significant changes to his 
monograph. I believe the work I have done herein is at least a strong advance 
forward toward completing a revision, and this is a “big-picture” goal I have 
considered ever since I began consulting his publication for assistance in 
identifying my field samples of echinoids. 

The goal of a dissertation is to learn how to ask appropriate questions, 
learn to solve any such questions, and to improve one’s personal knowledge 
while contributing to the science as a whole. I may not know everything about 
echinoids or their relatives, but I do know that the more I learn about them, the 
greater my curiosity grows to learn more. 


APPENDIX 

ECHINOID BIOMETRIC TRAIT MEASUREMENTS 


# 

UF , USNM # 

TL 

TW 

PSL 

PSW 

PSP 

PPL 

PPW 

1 

28204.01 

75.66 

75.60 

3.17 

2.87 

32.10 

2.37 

1.00 

2 

28204,02 

45.55 

44.98 

3.13 

2.87 

19.55 

1.39 

1.09 

3 

28204.03 

34.23 

33.92 

2.40 

2.13 

14.10 

• 

• 

4 

28204.04 

33.38 

32.67 

1.80 

1.66 

13.98 

1.37 

0.91 

5 

28204.05 

25.64 

27,67 

1.90 

1.85 

10.87 

1.42 

0.83 

6 

28204.06 

21.79 

21.80 

1.29 

1.16 

9.29 

0.87 

0.48 

7 

30401.01 

72.21 

71.85 

2.76 

2,55 

31.72 

2.18 

1.28 

8 

30401.02 

44.45 

45.46 

2.84 

2.65 

18.30 

• 

• 

9 

30401.03 

42.79 

42.76 

1.66 

1.49 

19.15 

1.43 

0.86 

10 

30401.04 

43.18 

44,65 

2.45 

2.18 

18.36 

1.70 

1.13 

11 

30401.05 

38.15 

39.38 

2.50 

2.27 

15.57 

1.49 

1.06 

12 

30401.06 

34.62 

35.02 

1.80 

1.75 

14.13 

1.48 

0.81 

13 

30401.07 

30.14 

31.03 

1.80 

1.72 

12.59 

1.58 

0.91 

14 

30401.08 

27.64 

28.84 

1.67 

1.43 

11.68 

1.24 

0.78 

15 

30401.09 

20.72 

21.34 

1.32 

1.27 

8.55 

0.86 

0.60 

16 

30401.10 

• 

• 

1.52 

1.44 

• 

1.34 

0.68 

17 

28207.01 

74,34 

73.83 

3.04 

2.94 

32.43 

1.63 

1.14 

18 

28207.02 

64.52 

65.04 

2.48 

2.24 

29.10 

1.54 

1.19 

19 

28207.03 

59,48 

• 

3.28 

3.04 

26.67 

2.36 

1.06 

20 

21435.01 

74,10 

75.99 

• 

• 

• 

• 

• 

21 

24518.01 

36.42 

37.70 

1.79 

1.63 

15.04 

1.70 

0.58 

22 

24518.02 

27.76 

27.54 

1.67 

1.62 

11.86 

0.96 

0.87 

23 

28208.01 

35.54 

34.87 

2.03 

1.73 

15.40 

1.28 

0.81 

24 

28208.02 

33.59 

34.03 

2.07 

1.91 

13.53 

1.42 

0,90 

25 

28208.03 

40.99 

• 

1.82 

1.79 

17.75 

1.71 

1.02 

26 

28208.04 

31.82 

32.66 

2.36 

2.10 

13.10 

1.24 

0.90 

27 

28208,05 

• 

33.59 

1.47 

1,42 

14.35 

1.72 

0.68 

28 

21312.01 

55.79 

56.80 

2.94 

2.73 

25.17 

2.18 

1.18 

29 

21312.02 

40.30 

40.04 

2.47 

2.33 

17.01 

1.20 

0.95 

30 

21312.03 

34.62 

• 

1.95 

1.80 

14.33 

1.42 

0.78 

31 

21312.04 

28.65 

29.14 

1.32 

1.28 

12.35 

1.40 

0.63 

32 

21312.05 

23.27 

23.68 

1.57 

1.73 

9.98 

1.02 

0.85 

33 

21312.06 

20.06 

20.16 

1.29 

1.38 

8.50 

0.96 

0.60 

34 

21312.07 

18,83 

19.25 

1.16 

1.39 

7.92 

0.99 

0.67 

35 

21313.01 

64.51 

63.51 

2.56 

2.43 

27.66 

1.85 

1.15 

36 

21313.02 

58.29 

58.31 

2.66 

2.55 

25.60 

1.48 

0.91 

37 

21313.03 

65.03 

65.87 

2,78 

2.67 

28.57 

1,75 

1.19 

38 

21313.04 

33.37 

34.49 

1.48 

1.49 

13.89 

1.58 

0,77 

39 

21313.05 

31.54 

32.76 

2.00 

1.82 

13.43 

1.05 

0.80 

40 

21313.06 

30.37 

30.30 

1.58 

1.47 

12.92 

1.32 

0.80 

41 

21313,07 

30.94 

30.35 

1.48 

1.34 

13.29 

1.04 

0.68 

42 

21313.08 

28.63 

28.32 

2.34 

2.15 

11.84 

0.99 

0.91 

43 

21313.09 

28.66 

28.63 

1.49 

1.40 

11.92 

1.38 

0.62 

44 

21313.10 

26.41 

27.10 

1.66 

1.48 

11.48 

1.30 

0.68 

45 

21313.11 

21.33 

21.38 

1.63 

1,48 

8,20 

1.18 

0.82 

46 

21443.01 

77.49 

76.82 

• 

• 

35.10 

• 

• 

47 

21443.02 

• 

30.93 

1,80 

1.86 

• 

1.06 

0.93 

48 

21443.03 

25,40 

25,38 

1.65 

1.57 

10.50 

0.93 

0.91 


357 


# 

49 

50 

51 

52 

53 

54 

55 

56 

57 

58 

59 

60 

61 

62 

63 

64 

65 

66 

67 

68 

69 

70 

71 

72 

73 

74 

75 

76 

77 

78 

79 

80 

81 

82 

83 

84 

85 

86 

87 

88 

89 

90 

91 

92 

93 

94 

95 

96 


358 


UF , USNM # 

TL 

TW 

PSL 

PSW 

PSP 

PPL 

PPW 

21443.04 

24.89 

26.30 

1.19 

1.24 

10.78 

0.83 

0.48 

21443.05 

25.04 

25.90 

1.06 

0.97 

10.31 

1.37 

0.50 

21443.06 

22.96 

22.74 

1.02 

1,04 

9.62 

1.23 

0.58 

21443.07 

22.00 

23.62 

1.23 

1.14 

9.29 

0.83 

0.44 

21443.08 

19.38 

19.54 

1.21 

0.91 

9.05 

0,62 

0.44 

21443.09 

19.53 

20.75 

1.09 

1.09 

8.50 

1.06 

0.57 

21443.10 

18.66 

18.69 

0.99 

0.97 

7.98 

0.86 

0.48 

21443.11 

16.63 

17.64 

1.09 

1.18 

7.06 

0.82 

0.45 

21443.12 

14.45 

14.73 

1.07 

1.01 

6.04 

0,71 

0.26 

40360.01 

43.59 

42.32 

2.23 

2,13 

19.15 

1.60 

1.24 

40362.01 

41.41 

42.18 

1.82 

1.65 

18.07 

1.96 

0.72 

40363.01 

58.57 

58.26 

2.71 

2.74 

25.80 

1.85 

1.15 

40364.01 

38.64 

38.49 

2.27 

2.13 

16,82 

1.89 

0.80 

40385.01 

32.18 

32.61 

1.54 

1.49 

14.30 

1.32 

0.76 

40386.01 

44.05 

44.74 

2.31 

2.18 

18.05 

2.15 

0.73 

40387.01 

32.82 

32.62 

1.73 

1,84 

13.89 

1,07 

0.92 

40366.01 

34.15 

33.27 

2.04 

1.93 

14.84 

1.46 

0.74 

40367.01 

76.70 

76.16 

3.12 

3,11 

33.60 

1.96 

1.15 

40421.01 

36.96 

35.17 

1.62 

1.67 

15.12 

1,79 

0.66 

40422.01 

42.29 

41.60 

2.62 

2.43 

18.07 

2.00 

1.07 

40371.01 

53.61 

52.92 

3,25 

2,89 

22.60 

2.31 

1.14 

40372.01 

• 

34.18 

1.67 

1.42 

13.94 

1.94 

0.63 

40373.01 

40.47 

39.44 

2,26 

2,26 

17,17 

1.38 

0,92 

40374.01 

33.38 

32.86 

2.13 

1.82 

13.88 

2.18 

1,01 

40370.01 

• 

61.68 

2.69 

2.45 

28.61 

• 

• 

40375.01 

36.39 

37.54 

2,12 

2.03 

15.36 

1.61 

0.87 

40389.01 

36.22 

37.38 

1.54 

1.51 

15.60 

1.60 

0.83 

40368.01 

43.24 

• 

2.04 

1.84 

18.93 

1.63 

0.77 

40431.01 

79.34 

80.44 

3.22 

3.25 

35.59 

• 

• 

40380.01 

27.86 

28.27 

1.62 

1.44 

11.98 

1.52 

0.59 

40379.01 

30.44 

30.49 

2.10 

2.13 

12.70 

1.23 

0.83 

40365.01 

26.98 

28.00 

1.81 

1.75 

11.41 

1.06 

0.63 

40358.01 

26.61 

26.72 

1.23 

1.21 

11.40 

1.02 

0.67 

40359.01 

28.89 

28.63 

• 

• 

12.02 

1.18 

0.78 

40388.01 

24.09 

24.98 

1.44 

1.20 

10.32 

• 

• 

40369.01 

28.75 

29.01 

1.52 

1.62 

12,38 

1.25 

0.74 

40417.01 

42.84 

• 

1.48 

1.65 

19.20 

2.38 

0.69 

40418.01 

35.19 

• 

1.70 

1.67 

14.42 

1.16 

0,76 

40419.01 

30.75 

32.02 

1.99 

2.00 

12.70 

1.20 

0.86 

40420.01 

42.13 

42.17 

1.90 

1.90 

18,04 

1.96 

0.68 

40432.01 

78.93 

78.91 

2.89 

3.08 

34.31 

• 

• 

40384.01 

25.47 

• 

1.60 

1.49 

10.84 

1.51 

0.76 

40383.01 

29.01 

29.54 

1.43 

1.57 

12.66 

1.25 

0.71 

40381.01 

27.41 

27.76 

1.47 

1.33 

11.88 

1.80 

0.60 

40391.01 

25.42 

27.03 

1.11 

1.20 

11.37 

1.27 

0.60 

40390.01 

27.00 

26.79 

1.62 

1.56 

11,43 

1.53 

0.86 

40377.01 

27.12 

26.23 

• 

1,56 

11.11 

1.27 

0,66 

40376.01 

28.47 

28.00 

1.54 

1.27 

12.11 

1.42 

0.63 


359 


# 

UF , USNM # 

TL 

TW 

PSL 

PSW 

PSP 

PPL 

PPW 

97 

40361.01 

25.45 

26.17 

1.21 

1.21 

11.18 

1.23 

0.53 

98 

40429,01 

80.03 

79.59 

3.04 

2.79 

36.37 

1.98 

1.32 

99 

40430.01 

59.75 

60.24 

2.54 

2.36 

25.90 

2.00 

0.99 

100 

40428.01 

70.42 

70.14 

3.02 

2.95 

30.55 

2.18 

1.39 

101 

40433.01 

54,67 

55.39 

2.36 

2.27 

23,87 

1,99 

0.96 

102 

29684.01 

64.68 

63.61 

2.64 

2.59 

29.19 

1.86 

0.88 

103 

29684.02 

30.06 

30.42 

2.13 

2.00 

12.57 

1.14 

0.77 

104 

29684.03 

27.64 

26.53 

1.27 

1.20 

11.72 

1.20 

0.69 

105 

40416.01 

25.69 

27.02 

1,68 

1.68 

10.71 

1.15 

0.71 

106 

40415.01 

26.31 

27.20 

1.01 

• 

• 

1.01 

0.71 

107 

40414.01 

28.67 

29.65 

1.57 

1.53 

12.11 

1.25 

0.67 

108 

40413.01 

26.23 

27.66 

1.52 

1.56 

10.92 

0.96 

0.67 

109 

40411.01 

25.60 

25.85 

1.09 

1.09 

10.65 

0.97 

0.66 

110 

40409.01 

31.40 

30.91 

1.81 

1.71 

13.39 

1,49 

0.83 

111 

40382.01 

26.49 

27.50 

1.79 

1.73 

11.13 

1.34 

0.71 

112 

40434.01 

22.41 

22.19 

1.33 

1,34 

9.77 

1.14 

0.68 

113 

40426.01 

22.59 

23.11 

1.19 

1.11 

9,56 

0,96 

0.58 

114 

40425.01 

29.57 

29.38 

1.79 

1.76 

12.16 

1.11 

0.78 

115 

40424.01 

21.50 

21.64 

1.09 

1.14 

8.92 

0.86 

0.62 

116 

40423,01 

28.18 

28.11 

1.79 

1.77 

11.91 

1.27 

0.97 

117 

40412.01 

20.72 

20.68 

1.21 

1.11 

8.99 

1.01 

0.53 

118 

40392.01 

18.45 

18.40 

1.14 

1.06 

7.65 

0.91 

0.43 

119 

40378.01 

18.33 

17.78 

1.18 

1.13 

7.79 

0.83 

0.59 

120 

13753.01 

62.86 

66.15 

2.73 

2.76 

26.84 

1.91 

1.27 

121 

13753.02 

48.72 

48.97 

2,29 

2.18 

21.14 

1,47 

0.77 

122 

13753.03 

27.68 

28.04 

1.32 

1.40 

11.51 

1.16 

0.82 

123 

13753.04 

24,56 

25.31 

1.28 

1,25 

10.40 

0,90 

0.45 

124 

13753.05 

22.97 

22.64 

1.30 

1.44 

9.18 

0.96 

0.68 

125 

13753.06 

23.48 

23.20 

1.11 

1.20 

9.77 

0.90 

0.52 

126 

13753.07 

19.35 

19.34 

1.10 

1.05 

8.07 

0.92 

0.34 

127 

13075.01 

83.08 

83.88 

• 

• 

• 

• 

• 

128 

29670.01 

36.75 

38.13 

1.66 

1.58 

15.35 

1.96 

0.77 

129 

13076.01 

76.86 

76,70 

2.70 

2.73 

34.02 

• 

• 

130 

465436.01 

105.96 

103.72 

• 

• 

• 

• 

• 

131 

465463.01 

60.52 

59.09 

2.56 

2.26 

26.63 

2,46 

1.01 

132 

465464.01 

71.95 

70.00 

• 

• 

32.38 

4.80 

1.47 

133 

465449.01 

57.91 

57.92 

3.04 

2.70 

26.06 

2.57 

1.27 

134 

465440.01 

56.99 

56.19 

2.00 

1,72 

25,59 

2.66 

0.92 

135 

465458.01 

59.98 

58.35 

3.14 

3.28 

27.38 

1.87 

1.42 

136 

465467.01 

62,40 

62.38 

• 

• 

• 

• 

• 

137 

465444.01 

22.47 

22.80 

1.16 

1.16 

9.84 

0.91 

0.57 

138 

465466.01 

69.88 

68.68 

3.53 

3.65 

31.29 

2.17 

0.96 

139 

465451.01 

65.12 

63.24 

2.81 

2.66 

29.37 

2.01 

1,20 

140 

465465,01 

55,49 

55.29 

2.81 

2.66 

24.28 

3.46 

0,97 

141 

465456.01 

21.64 

21.03 

1.77 

2.01 

8.53 

1.38 

0.57 

142 

465448.01 

65.64 

65,04 

2.32 

2,59 

30.22 

1.70 

1.11 

143 

465445.01 

• 

126.96 

• 

• 

• 

• 

• 

144 

465443.01 

25.47 

25.40 

1.33 

1.28 

10.87 

0.83 

0.60 


# 

145 

146 

147 

148 

149 

150 

151 

152 

153 

154 

155 

156 

157 

158 

159 

160 

161 

162 

163 

164 

165 

166 

167 

168 

169 

170 

171 

172 

173 

174 

175 

176 

177 

178 

179 

180 

181 

182 

183 

184 

185 

186 

187 

188 

189 

190 

191 

192 


360 


UF , USNM # 

TL 

TW 

PSL 

PSW 

PSP 

PPL 

PPW 

465442.01 

31.40 

31.59 

1.56 

1.48 

13.42 

1.02 

0.66 

465468.01 

28.82 

29.69 

1.42 

1.33 

12.71 

1.11 

0.97 

438118.01 

70.56 

72.72 

• 

• 

• 

• 

• 

438188.02 

67.50 

65.18 

• 

• 

• 

2.87 

1.11 

1.01 

107.04 

114.49 

3.16 

2.98 

48.43 

5.35 

1.35 

2.01 

101.42 

103.73 

3.74 

3.68 

46.65 

5.37 

1.33 

3.01 

96.60 

98.24 

3.40 

3.14 

42.30 

5.16 

1.47 

5.01 

118.42 

121.81 

4.15 

3.71 

53.19 

3.93 

1.63 

9.01 

82.20 

84.88 

2.80 

3.03 

35.59 

2.93 

1.31 

10.01 

102.78 

104.44 

3.53 

3.67 

46.01 

5.23 

0.99 

11.01 

116.09 

120.08 

3.67 

3.45 

53.27 

4.51 

1.52 

12.01 

76.57 

76.15 

• 

• 

33.40 

• 

• 

16.01 

95.47 

• 

3.40 

3.13 

42.77 

4.92 

1.07 

17.01 

101.89 

106.87 

3.14 

2.87 

45.97 

5.81 

1.38 

19.01 

83.64 

89.30 

3.31 

3.21 

38.81 

4.76 

1.25 

20.01 

115.61 

120.02 

4.03 

3.56 

53.74 

3.25 

1.31 

21.01 

106.29 

111.01 

3.50 

3.50 

48.86 

7.08 

1.10 

22.01 

94.38 

95.70 

2.99 

2.94 

42.42 

3.10 

1.20 

23.01 

83.52 

86.30 

3.22 

3.13 

37.17 

2.82 

1.36 

24.01 

91.19 

93.61 

3.60 

3.26 

41.31 

5.20 

1.18 

25.01 

113.34 

116.31 

• 

2.55 

50.10 

3.25 

1.66 

26.01 

125.05 

• 

3.85 

3.72 

54.85 

2.49 

1.42 

27.01 

95.73 

96.85 

3.03 

2.95 

42.98 

6.97 

1.19 

28.01 

87.03 

• 

3.34 

3.11 

38.67 

4.10 

1.19 

32.01 

91.03 

91.05 

3.56 

3.13 

41.84 

4.61 

1.13 

33.01 

• 

72.35 

3.52 

3.32 

39.98 

4.97 

1.51 

35.01 

81.41 

84.22 

2.87 

2.91 

36.38 

2.88 

1.15 

36.01 

111.34 

117.41 

3.55 

3.44 

48.91 

2.49 

1.34 

37.01 

103.33 

111.49 

3.79 

3.54 

48.51 

4.14 

1.01 

38.01 

• 

93.24 

3.13 

3.01 

41.60 

3.88 

1.72 

39.01 

79.69 

79.24 

3.04 

2.85 

35.39 

4.07 

1.06 

44.01 

104.08 

110.66 

3.32 

3.17 

49.86 

3.23 

0.98 

46.01 

73.99 

76.10 

3.05 

2.93 

33.19 

2.49 

1.31 

49.01 

96.84 

• 

3.31 

3.21 

44.56 

5.28 

1.20 

51.01 

96.93 

101.51 

3.14 

3.19 

43.41 

4.58 

1.56 

55.01 

103.18 

108.16 

2.93 

3.27 

46.02 

6.04 

1.16 

58.01 

86.71 

• 

2.91 

2.58 

40.77 

3.87 

0.99 

69.01 

95.73 

95.01 

3.14 

2.97 

43.25 

2.74 

1.43 

70.01 

101.10 

104.53 

3.34 

3.20 

45.42 

3.10 

1.25 

72.01 

90.87 

93.29 

3.30 

3.19 

41.19 

2.29 

1.09 

77.01 

111.10 

113.54 

3.45 

3.54 

53.18 

3.52 

1.34 

79.01 

88.22 

88.77 

3.41 

3.15 

40.07 

5.37 

1.21 

81.01 

100.82 

103.68 

3.21 

3.10 

47.32 

2.01 

1.05 

89.01 

112.41 

115.17 

3.45 

3.17 

52.56 

2.44 

1.22 

96.01 

99.30 

104.63 

3.04 

2.93 

46.74 

4.57 

1.24 

97.01 

95.72 

100.12 

3.67 

3.23 

42.14 

2.60 

• 

98.01 

93.38 

97.25 

2.98 

2.73 

41.49 

5.88 

1.48 

99.01 

88.40 

91.14 

3.49 

3.36 

41.09 

3.61 

1.19 


# 

193 

194 

195 

196 

197 

198 

199 

200 

201 

202 

203 

204 

205 

206 

207 

208 

209 

210 

211 

212 

213 

214 

215 

216 

217 

218 

219 

220 

221 

222 

223 

224 

225 

226 

227 

228 

229 

230 

231 

232 

233 

234 

235 

236 

237 

238 

239 

240 


361 


UF, USNM# 

TL 

TW 

PSL 

PSW 

PSP 

PPL 

PPW 

105.01 

99.87 

105.44 

3.61 

3.18 

47.68 

5.32 

1.38 

108.01 

66.85 

68.96 

2.93 

2.94 

30.30 

3.61 

1.06 

109.01 

98.71 

101.74 

3.70 

3.59 

43.11 

6.88 

1.32 

116.01 

81.61 

84.44 

3.08 

3.04 

37.33 

5.49 

1.34 

118.01 

105.13 

106.93 

3.46 

3.31 

46.72 

5.43 

1.34 

119.01 

87.07 

95.60 

3.28 

3.08 

38.96 

4.27 

1.32 

124.01 

92.90 

96.48 

2.95 

3.06 

41.50 

5.90 

1.29 

127.01 

91.82 

98.31 

3.17 

3.16 

41.94 

5.84 

1.37 

129.01 

93.86 

95.90 

3.38 

3.08 

42.71 

2.15 

1.20 

141.01 

96.15 

105.77 

3.65 

3.69 

46.03 

5.27 

1.06 

143.01 

72.64 

74.72 

2.95 

2.74 

33.66 

3.62 

1.33 

144.01 

118.91 

123.97 

4.06 

3.94 

52.31 

3.36 

1.16 

146.01 

67.18 

68.70 

2.89 

2.62 

31.57 

2.99 

1.05 

147.01 

130.27 

134.54 

3.78 

3.70 

59.50 

5.21 

1.51 

150.01 

91.27 

92.72 

3.07 

2.95 

41.30 

4.07 

1.33 

169.01 

88.14 

90.78 

3.31 

3.30 

37.97 

4.60 

1.54 

171.01 

61.50 

• 

2.63 

2.47 

27.21 

2.36 

0.97 

174.01 

71.99 

72.54 

2.76 

2.65 

31.75 

2.06 

1.09 

184.01 

93.92 

97.87 

3.56 

3.53 

41.60 

1.95 

1.09 

185.01 

92.10 

93.13 

3.12 

3.10 

41.76 

3.45 

1.37 

194.01 

100.73 

102.27 

3.31 

3.18 

44.98 

3.60 

1.31 

201.01 

102.76 

103.82 

3.18 

3.40 

45.86 

4.32 

1.44 

202.01 

94.45 

97.14 

3.46 

3.37 

43.97 

3.42 

1.27 

204.01 

• 

107.14 

3.77 

3.54 

45.67 

3.03 

1.27 

207.01 

100.71 

105.22 

3.56 

3.34 

42.48 

2.22 

0.96 

208.01 

100.04 

106.86 

3.86 

3.46 

47.01 

2.50 

1.33 

209.01 

100.94 

102.94 

3.33 

3.03 

44.62 

3.25 

1.43 

210.01 

90.18 

91.89 

3.24 

3.18 

40.05 

3.17 

1.44 

211.01 

90.78 

93.64 

3.35 

3.27 

40.15 

2.23 

1.38 

212.01 

110.13 

• 

3.47 

3.25 

47.76 

4.42 

1.32 

465472.01 

100.83 

106.95 

• 

• 

• 

• 

• 

146726.01 

118.35 

• 

• 

• 

• 

5.76 

1.98 

465470.01 

101.57 

109.23 

• 

• 

• 

• 

• 

465470.02 

64.92 

68.70 

4.06 

3.12 

28.05 

3.78 

1.21 

154281.01 

151.30 

• 

5.23 

4.96 

69.10 

5.47 

1.57 

146720.01 

128.73 

139.34 

5.21 

4.30 

56.66 

5.79 

1.63 

146644.01 

119.84 

128.48 

5.08 

4.41 

51.40 

• 

1.27 

465471.01 

100.46 

104.92 

3.70 

4.01 

42.68 

3.91 

1.07 

146724.01 

• 

131.95 

5.46 

5.20 

50.83 

4.07 

1.67 

465469.01 

93.09 

95.51 

3.47 

3.45 

40.33 

3.36 

0.99 

21324.01 

90.69 

84.22 

3.94 

3.65 

39.01 

3.03 

1.67 

21324.02 

91.49 

91.31 

• 

4.48 

• 

• 

2.20 

21324.03 

94.19 

• 

3.84 

3.64 

38.84 

2.99 

2.08 

21324.04 

87.35 

80.22 

4.81 

4.85 

34.82 

2.48 

2.28 

21324.05 

83.45 

78.90 

3.91 

3.63 

35.50 

2.47 

1.32 

21324.06 

70.10 

65.41 

3.27 

3.14 

28.24 

2.38 

1.34 

21324.07 

56.93 

55.84 

3.02 

2.99 

24.20 

2.33 

1.20 

24511.01 

74.02 

71.19 

4.29 

4.34 

30.84 

2.64 

1.57 


362 


# 

UF , USNM # 

TL 

TW 

PSL 

PSW 

PSP 

PPL 

PPW 

241 

24511.02 

59.76 

55.65 

3.70 

3.58 

24.51 

1.76 

1.18 

242 

24511.03 

54.55 

47.98 

3.47 

3.40 

20.32 

1.57 

1.42 

243 

24511.04 

47.61 

43.87 

• 

• 

• 

1.94 

1.29 

244 

24511.05 

45.16 

42.90 

3.32 

3.07 

17.98 

1.93 

1.20 

245 

24511.06 

42.73 

40.01 

2.98 

3.00 

17.61 

1.32 

1.11 

246 

24511.07 

43.28 

40.00 

2.79 

2.76 

16.71 

1.43 

1.06 

247 

24511.08 

40.65 

37.45 

2.80 

2.90 

16.30 

1.90 

1.20 

248 

24511.09 

41.00 

36.46 

3.13 

3.18 

16.59 

1.99 

0.96 

249 

24511.10 

39.58 

35.89 

2.83 

2.90 

15.62 

1.25 

1.05 

250 

24511.11 

38.58 

36.37 

2.71 

2.80 

14.75 

1.28 

1.06 

251 

24511.12 

39.12 

35.10 

3.17 

3.08 

15.68 

1.67 

1.11 

252 

24511.13 

36.28 

33.33 

2.67 

2.54 

14.36 

1.80 

1.05 

253 

24511.14 

37.04 

33.94 

2.62 

2.79 

14.49 

1.32 

0.95 

254 

24511.15 

25.88 

24.96 

2.01 

1.96 

10.32 

1.23 

0.74 

255 

24511.16 

35.80 

33.95 

2.41 

2.70 

14.03 

1.38 

1.09 

256 

24511.17 

32.48 

30.64 

2.71 

2.71 

13.22 

1.09 

1.06 

257 

24511.18 

33.33 

30.73 

2.83 

2.67 

13.77 

1.20 

1.00 

258 

24511.19 

33.42 

29.28 

2.69 

2.61 

13.13 

1.13 

0.96 

259 

24511.20 

30.22 

28.49 

2.57 

2.23 

11.73 

1.48 

0.93 

260 

24511.21 

29.81 

28.21 

2.05 

2.18 

12.50 

1.10 

0.85 

261 

24511.22 

29.66 

26.80 

2.09 

2.10 

11.83 

1.52 

0.91 

262 

24511.23 

21.90 

20.42 

1.79 

1.85 

8.83 

1.09 

0.81 

263 

21212.01 

68.55 

67.53 

3.61 

3.80 

27.43 

2.51 

1.78 

264 

21213.01 

74.12 

69.62 

3.57 

3.95 

29.93 

1.99 

1.62 

265 

21214.01 

54.58 

51.10 

3.40 

3.24 

22.61 

1.79 

1.28 

266 

21215.01 

100.44 

98.03 

• 

• 

• 

3.67 

2.30 

267 

21216.01 

100.97 

95.97 

4.07 

4.10 

42.35 

3.57 

1.85 

268 

21217.01 

109.52 

106.24 

4.73 

5.05 

46.58 

3.87 

2.39 

269 

21218.01 

102.94 

99.01 

5.71 

6.03 

41.37 

4.54 

1.95 

270 

21219.01 

94.64 

96.65 

4.18 

3.92 

39.17 

3.03 

1.47 

271 

21220.01 

91.65 

92.75 

• 

• 

• 

3.04 

1.68 

272 

21221.01 

76.34 

70.14 

3.73 

3.96 

31.30 

2.50 

2.27 

273 

21223.01 

70.70 

64.08 

3.71 

4.10 

28.92 

2.46 

1.64 

274 

21224.01 

62.11 

60.85 

3.19 

3.36 

25.09 

2.44 

1.31 

275 

21225.01 

66.03 

63.76 

3.54 

3.65 

27.60 

2.11 

1.66 

276 

21226.01 

43.78 

41.50 

2.85 

2.84 

17.26 

1.86 

1.51 

277 

21227.01 

68.12 

66.27 

4.13 

4.05 

28.97 

3.18 

1.66 

278 

21228.01 

62.49 

59.39 

3.63 

3.52 

25.43 

2.12 

1.62 

279 

21229.01 

49.73 

43.19 

3.19 

3.41 

19.18 

1.84 

1.30 

280 

21230.01 

69.20 

64.14 

4.20 

4.27 

27.07 

2.94 

1.70 

281 

21231.01 

55.10 

52.65 

3.53 

3.60 

22.30 

2.42 

1.52 

282 

21232.01 

68.17 

65.02 

3.70 

3.85 

27.38 

3.01 

1.31 

283 

21233.01 

66.33 

63.60 

3.68 

3.61 

27.09 

2.02 

1.56 

284 

21234.01 

62.98 

60.26 

3.10 

2.99 

26.89 

1.75 

1.23 

285 

21235.01 

48.64 

44.31 

3.37 

3.34 

20.36 

1.66 

1.38 

286 

21236.01 

62.77 

62.87 

3.46 

3.65 

24.75 

2.26 

1.49 

287 

21237.01 

57.10 

50.66 

3.25 

3.06 

23.12 

1.93 

1.44 

288 

21238.01 

50.42 

47.38 

• 

• 

21.04 

1.81 

1.39 


# 

289 

290 

291 

292 

293 

294 

295 

296 

297 

298 

299 

300 

301 

302 

303 

304 

305 

306 

307 

308 

309 

310 

311 

312 

313 

314 

315 

316 

317 

318 

319 

320 

321 

322 

323 

324 

325 

326 

327 

328 

329 

330 

331 

332 

333 

334 

335 

336 


363 


UF , USNM # 

TL 

TW 

PSL 

PSW 

PSP 

PPL 

PPW 

21239.01 

48.83 

45.67 

3.07 

2.99 

19.49 

1.54 

1.24 

21240.01 

57.94 

55.96 

4.69 

4.60 

21.93 

2.16 

1.53 

21241.01 

49.44 

45.30 

3.22 

3.16 

20,11 

1.53 

1.13 

21242.01 

42.62 

41.24 

• 

• 

• 

• 

1.13 

21243.01 

48.51 

44.18 

3.14 

3.15 

18.17 

2.10 

1.54 

21244.01 

44.53 

40.92 

2.93 

3.10 

18.14 

1.57 

1.23 

21245.01 

51.96 

49.75 

3.07 

3.11 

21.09 

2.03 

1.36 

21246.01 

45.32 

42.65 

3.13 

3.05 

19.32 

1.65 

1.17 

21247.01 

41.65 

38.62 

2.83 

2.61 

16.67 

1.59 

1.32 

21248.01 

44.06 

41.25 

2.89 

3.32 

18.34 

1.70 

1.23 

21249.01 

45,33 

42.48 

2.78 

3.02 

18.49 

1.34 

1.12 

21250.01 

38.38 

37.58 

2.65 

2.54 

15.89 

1.63 

1.12 

21251.01 

39.74 

38.81 

2,59 

2.65 

16.32 

2.06 

1.07 

21252.01 

36.78 

35.51 

2.70 

2,65 

15.04 

1.19 

0.99 

21253.01 

39.06 

37.01 

2.59 

2,74 

15.89 

1.53 

1.13 

21254.01 

32.87 

30.80 

2.50 

2.51 

13.11 

1.40 

0.90 

21255.01 

35.59 

33.95 

2.60 

2.82 

13.89 

1.20 

0.95 

21256.01 

34.20 

31.69 

2.51 

2.62 

13.83 

1.57 

0.88 

21257.01 

35.50 

35.16 

1.79 

1.80 

15.75 

1.37 

0.85 

21258.01 

34.90 

33.21 

2.30 

2.26 

13.99 

2.16 

0.98 

21259.01 

38.94 

36.91 

2.70 

2.73 

16.01 

1.49 

1.44 

21260.01 

38.13 

35.26 

2.62 

2.43 

15.07 

1.38 

1.21 

21261.01 

36.37 

34.59 

2.40 

2.63 

14.43 

1.50 

1.21 

21262.01 

40.28 

37.97 

2.58 

2.56 

15.36 

1.31 

0.93 

21263.01 

40.46 

37.89 

2.71 

2.82 

16.86 

1,66 

1.22 

21264.01 

45.53 

42.75 

• 

2.94 

18.49 

1.32 

1.07 

21265.01 

32.54 

30.36 

2.28 

2.24 

13.21 

1.16 

0.92 

21266.01 

33.34 

30.32 

2.26 

2.40 

13.42 

1.76 

1.12 

21267.01 

39.26 

37.40 

2.81 

2.88 

15.15 

1.44 

1.14 

21268.01 

34.26 

32.89 

2.12 

2.37 

13.47 

1.11 

0.85 

21269.01 

31.52 

29.65 

2.26 

2.32 

12.84 

1.04 

0.86 

21270.01 

30.61 

28.48 

2.63 

2.31 

12.47 

1.85 

• 

21271.01 

30.06 

27.44 

2.52 

2.61 

12.07 

1.48 

1.12 

21272.01 

35.77 

34.14 

2.68 

2.80 

14.74 

1.39 

1.09 

21273.01 

28.03 

26.16 

1.97 

2.02 

11.17 

1.33 

0.85 

21274.01 

30.98 

29,29 

2.25 

2.24 

12.33 

1.47 

0,95 

21275.01 

25.98 

24.67 

1.79 

2.22 

10.48 

1.11 

0.85 

21276.01 

23.86 

22.86 

2.01 

1.98 

9.04 

1.28 

0.83 

21277.01 

24.26 

22.91 

1.81 

1.86 

9.88 

1.20 

0.84 

21278.01 

22.94 

21.57 

1.70 

1.83 

9.57 

1.11 

0,85 

21279.01 

23.20 

20.75 

1.94 

1.97 

9.31 

0.99 

0.77 

21280.01 

21.02 

20.29 

1.72 

1.73 

8.69 

1.20 

0.91 

21281.01 

21.53 

20.27 

1.75 

1.74 

9.04 

1.15 

0,84 

21282.01 

20.75 

19.08 

1.55 

1.60 

8.57 

1.24 

0.79 

21283.01 

20.16 

18.87 

1.67 

1.77 

8.48 

1.07 

0.81 

28219.01 

100.98 

97.54 

4.93 

4.78 

41.97 

2.37 

1.98 

28219.02 

70.34 

65.15 

3.71 

3.83 

29.86 

1.92 

1.71 

28219.03 

57.74 

57.08 

3.54 

3.49 

23.38 

2.24 

1.78 


# 

337 

338 

339 

340 

341 

342 

343 

344 

345 

346 

347 

348 

349 

350 

351 

352 

353 

354 

355 

356 

357 

358 

359 

360 

361 

362 

363 

364 

365 

366 

367 

368 

369 

370 

371 

372 


364 


UF , USNM # 

TL 

TW 

PSL 

PSW 

PSP 

PPL 

PPW 

28219.04 

55.98 

51.42 

3.36 

3.51 

22.68 

2.22 

1.59 

28219.05 

51.48 

49.41 

3.19 

3.11 

20.36 

1.68 

1.33 

28219.06 

49.11 

46.88 

3.39 

3.42 

18.27 

1.87 

1.30 

28219.07 

48.27 

44.98 

3.19 

3.22 

20.24 

1.78 

1.49 

28219.08 

47.07 

43.98 

3.02 

3.01 

19.02 

1.72 

1.26 

28219.09 

45.25 

46.52 

2.18 

2.43 

18.83 

1.63 

0.95 

28219.10 

43.86 

42.71 

2.58 

2.80 

17.79 

1.72 

1.24 

28219.11 

42.07 

38.18 

2.82 

2.79 

16.97 

1.54 

1.24 

28219.12 

39.63 

37.07 

2.50 

2.45 

16.32 

1.66 

1.10 

28219.13 

38.27 

35.64 

2.61 

2.68 

15.55 

1.38 

1.06 

28219.14 

33.68 

31.28 

2.54 

2.53 

13.33 

1.15 

0.88 

28219.15 

27.51 

25.73 

2.20 

2.16 

11.16 

1.57 

0.84 

28219.16 

26.40 

24.64 

2.06 

2.11 

10.73 

1.16 

0.88 

28219.17 

25.23 

24.65 

1.94 

2.03 

10.81 

0.85 

0.57 

28219.18 

24.08 

23.29 

1.72 

1.94 

10.01 

1.28 

0.87 

28219.19 

21.08 

19.68 

1.80 

1.80 

8.88 

1.03 

0.83 

28219.20 

16.76 

14.88 

1.64 

1.62 

6.65 

0.91 

0.67 

465487.01 

65.43 

66.66 

3.17 

3.08 

29.17 

2.22 

1.39 

465487.02 

87.02 

82.14 

2.45 

2.60 

39.52 

• 

• 

465487.03 

89.10 

88.78 

• 



• 

• 

465487.04 

86.51 

81.87 

• 

• 

• 

• 

• 

465487.05 

• 

• 

• 


• 

• 

• 

465487.06 

89.92 

89.20 

• 

• 


2.67 

1.77 

465487.07 

97.02 

92.63 

• 



• 

• 

465487.08 

90.24 

• 

• 



• 

• 

465486.01 

67.84 

65.86 

2.45 

2.37 

31.77 

2.22 

1.15 

465486.02 

63.39 

65.74 

• 

• 

• 

• 

• 

154280.01 

79.13 

69.83 

4.78 

4.66 

29.29 

• 

• 

2512.01 

66.97 

61.18 

• 

• 

• 

3.07 

1.99 

2512.02 

31.30 

28.53 

2.09 

2.08 

12.19 

1.42 

1.07 

2512.03 

28.62 

25.71 

2.40 

2.19 

10.37 

1.47 

0.96 

2512.04 

27.31 

24.77 

2.92 

3.04 

10.32 

• 

• 

2512.05 

26.14 

23.95 

2.24 

2.31 

10.13 

1.66 

1.00 

145411.01 

89.16 

77.25 


• 

33.64 

• 

• 

12900.01 

90.91 



• 

• 

1.24 

1.16 

12901.01 

143.29 

139.52 

5.01 

4.27 

66.26 

• 

• 


# 

1 

2 

3 

4 

5 

6 

7 

8 

9 

10 

11 

12 

13 

14 

15 

16 

17 

18 

19 

20 

21 

22 

23 

24 

25 

26 

27 

28 

29 

30 

31 

32 

33 

34 

35 

36 

37 

38 

39 

40 

41 

42 

43 

44 

45 

46 

47 

48 


365 


ppp 

POSAP 

ANLL 

7.31 

34.11 

17.86 

5.86 

21.32 

10.29 

4.45 

15.29 

8.50 

3.86 

15.43 

7.46 

3.50 

11.77 

5.28 

2.69 

9.98 

3.78 

5.67 

32.66 

16.53 

5.51 

19.96 

8.06 

3.92 

19.58 

8.61 

5.27 

19.11 

10.08 

5.05 

17.25 

8.53 

3.31 

15.58 

7.39 

3.68 

13.75 

4.49 

3.35 

12.39 

5.65 

2.48 

9.37 

3.89 

3.46 

• 

7.31 

7.01 

33.92 

20.25 

6.03 

31.29 

13.43 

6.57 

28.42 

12.72 

• 

34.30 

16.76 

3.49 

16.38 

6.56 

3.91 

12.39 

5.58 

4.11 

16.58 

6.21 

4.30 

15.20 

7.78 

3.74 

18.85 

8.47 

4.15 

14.88 

6.65 

3.09 

14.99 

6.18 

6.70 

26.72 

11.55 

5.05 

19.07 

8.42 

3.96 

15.11 

6.74 

2.83 

13.27 

5.00 

3.32 

10.57 

4.77 

2.92 

9.34 

3.50 

2.42 

8.38 

3.35 

6.70 

29.41 

15.16 

5.89 

27.63 

15.10 

6.45 

30.89 

15.20 

2.98 

14.90 

8.49 

4.12 

14.73 

5.49 

3.25 

13.90 

5.34 

3.00 

14.05 

6.27 

4.54 

13.38 

6.35 

3.08 

13.32 

7.17 

3.27 

13.04 

5.15 

3.17 

9.49 

4.17 

• 

• 

18.00 

3.84 

• 

6.52 

3.31 

11.51 

6.01 


ANLW 

ANLP 

PDI 

3.16 

10.26 

12.82 

3.06 

7.03 

8.03 

1.98 

5.44 

• 

1.98 

4.39 

5.34 

2.18 

4.59 

4.41 

1.38 

4.05 

• 

2.29 

10.19 

11.92 

2.42 

7.97 

8.14 

2.75 

6.18 

6.31 

3.35 

7.06 

8.34 

2.97 

6.07 

6.65 

1.90 

5.74 

6.60 

1.60 

5.76 

6.18 

1.95 

5.02 

5.85 

1.67 

3.67 

• 

1.77 

5.48 

5.61 

5.02 

8.12 

12.89 

2.62 

8.29 

11.22 

2.66 

9.66 

10.78 

3.60 

10.13 

12.72 

1.62 

6.09 

6.14 

1.98 

5.63 

5.34 

1.68 

6.07 

6.38 

2.46 

5.48 

6.22 

2.05 

5.89 

6.68 

2.46 

5.35 

6.27 

1.68 

5.49 

• 

3.58 

9.25 

10.56 

2.38 

5.88 

7.15 

1.90 

5.89 

6.45 

1.75 

4.77 

• 

2.18 

4.26 

4.64 

1.43 

3.88 

4.36 

1.46 

4.08 

3.87 

2.60 

8.71 

10.82 

2.97 

7.20 

10.32 

3.08 

8.57 

11.43 

1.71 

4.77 

5.10 

1.81 

5.43 

5.68 

1.57 

5.20 

• 

1.80 

4.62 

5.27 

2.14 

4.74 

5.58 

2.03 

4.17 

6.50 

1.99 

4.26 

4.87 

1.66 

4.07 

4.27 

4.41 

• 

12.14 

2.04 

5.42 

5.96 

2.20 

4.15 

4.24 


PDII 

PDIII 

PDIV 

13.52 

12.80 

13.41 

8.47 

8.14 

9.05 

6.69 

6.29 

5.90 

4.66 

• 

5.35 

13.22 

12.34 

13.80 

8.64 

7.01 

8.50 

6.69 

• 

6.80 

8.97 

7.23 

8.53 

7.55 

• 

• 

7.09 

6.33 

7.32 

5.96 

4.82 

6.13 

6.00 

5.61 

5.81 

6.17 

5.14 

5.16 

12.73 

11.65 

12.42 

11.45 

11.08 

12.01 

11.32 

9.82 

10.79 

5.89 

• 

6.01 

5.51 

5.13 

5.77 

6.95 

6.50 

7.16 

7.37 

5.53 

7.11 

• 

• 

6.28 

6.81 

6.26 

6.94 

6.61 

6.42 

6.56 

11.60 

9.56 

11.58 

7.78 

• 

7.23 

• 

5.91 

• 

4.53 

• 

4.82 

5.00 

• 

4.59 

• 

• 

4.27 

11.06 

10.09 

10.28 

9.76 

9.82 

9.90 

11.91 

10.03 

11.58 

5.91 

5.80 

6.43 

6.24 

• 

6.50 

5.47 

• 

• 

5.56 

• 

5.25 

6.04 

5.60 

6.29 

6.78 

• 

6.40 

5.06 

• 

5.75 

13.98 

13.16 

13.85 

5.79 

• 

5.89 

4.82 

4.59 

4.76 


366 


# 

PPP 

POSAP 

ANLL 

ANLW 

ANLP 

PDI 

PDII 

PDIII 

PDIV 

49 

2.94 

11.58 

5.02 

1.77 

4.14 

4.97 

4.67 

9 

5.13 

50 

2.56 

10.99 

5.52 

1.51 

4.61 

5.00 

4.74 

4.47 

4.87 

51 

2.51 

10.80 

4.11 

1.38 

3.69 

4.14 

4.68 

4.11 

3.67 

52 

2.56 

10.27 

4.14 

1.21 

3.61 

• 

• 

• 

• 

53 

2.41 

10.08 

3.16 

0.99 

3.40 

• 

• 

• 

• 

54 

2.59 

8.85 

2.79 

1.47 

3.87 

• 

• 

• 

• 

55 

2.18 

8.43 

3.73 

1.53 

3.28 

• 

• 

• 

• 

56 

2.29 

8.03 

3.27 

1.66 

3.09 

• 

• 

• 

« 

57 

2.15 

6.80 

2.01 

1.21 

2.92 

• 

• 

• 

• 

58 

4.50 

20.01 

9.19 

2.26 

7.12 

• 

8.61 

• 

8.31 

59 

3.60 

19.10 

8.25 

1.95 

6.29 

8.68 

9.34 

9.53 

9.39 

60 

5.48 

27.97 

13.70 

2.60 

8.17 

11.06 

10.90 

9.60 

11.16 

61 

4.78 

17.91 

7.36 

2.57 

6.36 

7.51 

7.48 

6.71 

7.81 

62 

3.69 

• 

6.19 

1.13 


6.32 

6.92 

6.62 

7.13 

63 

4.86 

20.04 

9.34 

2.38 

7.79 

7.72 

8.30 

• 

8.57 

64 

4.71 

14.64 

7.84 

2.19 

5.89 

6.88 

7.64 

6.22 

7.55 

65 

4.03 

15.87 

6.87 

2.56 

5.99 

6.99 

6.71 

• 

6.80 

66 

7.25 

35.89 

18.99 

3.40 

10.24 

13.20 

14.23 

12.66 

14.22 

67 

3.39 

16.76 

8.89 

2.10 

5.51 

6.32 

6.76 

6.01 

6.80 

68 

4.88 

18.91 

9.62 

2.07 

6.75 

7.67 

7.96 

6.47 

7.96 

69 

5.88 

24.68 

14.09 

3.30 

7.73 

10.23 

10.22 

8.78 

10.84 

70 

3.39 

15.07 

6.60 

1.72 

6.61 

6.59 

7.21 

6.47 

7.31 

71 

5.01 

19.58 

8.61 

2.65 

5.96 

7.59 

8.10 

7.22 

8.15 

72 

3.25 

15.06 

5.48 

1.53 

6.57 

6.12 

5.96 

• 

• 

73 


26.44 

13.43 

2.48 

7.41 

11.48 

10.89 

10.40 

11.16 

74 

4.27 

17.03 

7.36 

2.40 

6.33 


7.06 

• 

7.18 

75 

3.22 

16.99 

7.26 

2.45 

5.25 

5.94 

6.43 

• 

6.45 

76 

4.52 

19.97 

9.20 

2.28 

6.84 

7.06 

7.26 

7.01 

7.92 

77 

7.74 

36.32 

20.91 

2.67 

9.90 

13.18 

13.74 

14.16 

14.27 

78 

3.58 

13.06 

4.90 

1.87 

5.54 


• 

• 

• 

79 

4.14 

14.08 

7.11 

2.24 

5.20 

5.71 

5.84 

5.23 

6.24 

80 

3.77 

12.31 

5.34 

2.05 

5.13 

5.44 

5.91 

5.48 

6.47 

81 

2.85 

12.52 

4.30 

1.49 

4.11 

4.81 

5.14 

4.59 

4.97 

82 

3.17 

12.78 

5.88 

1.71 

5.19 

5.13 

• 

• 

• 

83 

• 

11.21 

3.92 

1.51 

4.35 

4.58 

4.97 

• 

4.81 

84 

3.92 

13.37 

5.57 

1.47 

4.83 

5.14 

5.34 

5.66 

5.88 

85 

3.14 

20.63 

8.24 

1.85 

6.19 

6.41 


6.52 

7.59 

86 

3.58 

15.29 

7.34 

1.72 

5.39 

6.62 

7.08 

6.60 

6.85 

87 

4.21 

14.22 

7.30 

2.32 

5.13 

5.89 

6.35 

5.57 

6.42 

88 

4.00 

18.84 

8.76 

2.56 

7.35 

6.98 

7.09 

6.78 

7.36 

89 

6.60 

36.65 

19.64 

2.54 

8.80 

13.25 

14.37 

13.08 

14.37 

90 

3.11 

11.73 

4.49 

1.80 

4.87 

5.18 

5.35 

5.46 

5.42 

91 

3.47 

13.14 

6.17 

2.28 

5.21 

5.10 

5.38 

4.80 

5.51 

92 

2.84 

12.00 

5.53 

1.70 

5.58 

5.10 

5.58 

• 

5.46 

93 

2.51 

12.03 

4.52 

1.44 

4.03 

4.16 

4.80 

5.02 

5.27 

94 

3.34 

12.16 

4.74 

1.48 

5.10 

4.90 

5.18 

4.69 

5.33 

95 

3.45 

12.12 

6.12 

2.31 

4.77 

5.33 

6.01 

• 

6.19 

96 

3.14 

12.66 

5.28 

1.52 

4.80 

5.04 

• 

• 

• 


367 


# 

PPP 

POSAP 

ANLL 

ANLW 

97 

2.59 

• 

4.49 

1.67 

98 

7.84 

37.42 

18.77 

2.42 

99 

5.70 

27.38 

14.07 

3.32 

100 

7.09 

32.56 

16.12 

2.70 

101 

5.39 

• 

10.74 

3.60 

102 

5.62 

30.40 

15.86 

2.28 

103 

4.34 

14.02 

7.23 

2.34 

104 

2.74 

12.34 

5.39 

1.62 

105 

3.56 

11.04 

5.15 

2.00 

106 

3.07 

11.08 

4.83 

1.63 

107 

3.37 

13.09 

6.18 

1.84 

108 

3.78 

12.61 

3.25 

1.57 

109 

2.60 

11.30 

5.00 

1.70 

110 

3.49 

14.26 

7.49 

2.03 

111 

3.65 

11.69 

4.73 

1.84 

112 

2.81 

10.40 

4.87 

1.39 

113 

2.64 

10.38 

4.24 

1.44 

114 

3.81 

• 

5.88 

1.53 

115 

2.40 

9.33 

4.08 

1.56 

116 

3.75 

12.52 

6.18 

2.48 

117 

2.43 

9.60 

3.60 

1.66 

118 

2.13 

8.10 

3.04 

1.23 

119 

2.46 

8.42 

2.98 

1.18 

120 

5.57 

29.07 

16.92 

3.21 

121 

4.81 

22.16 

11.06 

2.78 

122 

3.00 

11.70 

5.52 

2.09 

123 

2.52 

11.12 

4.64 

1.71 

124 

2.84 

9.71 

4.59 

2.18 

125 

2.75 

10.52 

4.35 

1.40 

126 

2.13 

8.55 

3.64 

1.38 

127 

• 

37.80 

22.66 

2.61 

128 

3.49 

16.95 

6.43 

1.81 

129 

• 

35.83 

16.78 

3.65 

130 

• 

48.09 

31.11 

3.02 

131 

5.81 

27.55 

9.42 

1.60 

132 

7.07 

34.73 

10.65 

1.91 

133 

5.70 

27.15 

9.48 

1.94 

134 

4.87 

26.84 

10.56 

1.48 

135 

5.94 

29.08 

11.93 

2.08 

136 

• 

29.40 

10.13 

1.71 

137 

2.70 

10.80 

4.36 

1.46 

138 

6.68 

32.33 

12.16 

2.33 

139 

6.62 

31.43 

10.56 

1.54 

140 

4.54 

25.33 

11.30 

1.73 

141 

3.47 

• 

2.56 

1.32 

142 

6.37 

31.39 

12.02 

2.23 

143 

• 

• 

23.97 

4.26 

144 

2.60 

12.06 

4.24 

1.51 


ANLP 

PDI 

PDII 

PDIII 

PDiV 

• 

4.82 

5.09 

4.24 

4.85 

11.12 

15.10 

15.72 

14.10 

15.59 

8.07 

10.02 

10.76 

9.90 

10.83 

9.85 

11.88 

12.28 

11.98 

12.58 

• 

9.28 

9.96 

9.70 

10.68 

7.77 

10.76 

10.87 

11.01 

11.41 

4.97 

5.65 

5.98 

5.63 

6.09 

4.35 

3.96 

4.73 

• 

4.31 

5.23 

5.81 

6.57 

• 

• 

5.30 

4.86 

5.02 

4.54 

5.13 

4.91 

5.57 

6.32 

• 

• 

5.32 

4.95 

5.18 

4.52 

4.87 

4.02 

4.19 

4.78 

5.18 

4.54 

5.18 

5.62 

6.17 

5.72 

6.01 

5.65 

5.01 

5.13 

• 

5.04 

3.91 

4.29 

4.47 

• 

3.98 

4.15 

3.67 

3.96 

3.55 

3.88 

• 

5.19 

5.38 

5.10 

5.82 

3.72 

• 

4.30 

3.74 

4.48 

5.44 

5.52 

5.58 

4.91 

5.48 

3.93 

3.84 

4.16 

3.55 

• 

3.79 

3.54 

3.68 

• 

• 

3.77 

3.53 

3.77 

• 

3.58 

8.54 

11.59 

11.18 

10.55 

11.12 

6.52 

8.26 

8.01 

8.15 

8.75 

5.51 

5.02 

5.42 

5.70 

5.57 

3.84 

4.53 

4.22 

• 

4.58 

4.11 

• 

• 

• 

• 

3.89 

• 

• 

• 

• 

3.59 

3.72 

3.81 

3.55 

4.01 

11.16 

14.22 

15.06 

14.66 

15.43 

6.50 

6.36 

6.93 

• 

7.54 

10.24 

11.16 

12.82 

12.14 

13.04 

13.81 

• 

• 

• 

• 

13.97 

• 

• 

• 

• 

14.05 

10.28 

12.09 

10.54 

11.67 

11.65 

9.24 

10.64 

9.28 

7.35 

• 

8.66 

12.40 

12.52 

12.52 

11.07 

9.65 

10.52 

9.95 

10.65 

11.62 

9.69 

• 

• 

10.46 

3.94 

3.42 

3.63 

• 

3.91 

13.80 

11.15 

12.09 

11.15 

11.73 

12.62 

9.60 

10.38 

9.25 

10.62 

11.86 

9.20 

10.61 

8.85 

10.28 

• 

4.02 

• 

• 

• 

12.47 

10.09 

11.16 

10.89 

11.83 

• 

18.00 

18.05 

16.12 

20.21 

3.87 

• 

• 

• 

• 


368 


# 

PPP 

POSAP 

ANLL 

ANLW 

ANLP 

PDI 

PDII 

PDIII 

PDIV 

145 

3.37 

14.99 

6.00 

1.93 

4.54 

5.28 

5.53 

• 

5.43 

146 

3.81 

• 

3.82 

1.93 

• 

• 

• 

• 

• 

147 

• 

34.02 

14.21 

3.09 

14.10 

11.98 

12.75 

10.87 

12.70 

148 

• 

31.29 

11.88 

2.26 

13.61 

• 

11.56 

8.25 

10.73 

149 

6.41 

49.51 

13.81 

3.51 

21.83 

18.91 

18.97 

14.23 

20.51 

150 

7.77 

47.87 

15.18 

3.17 

19.58 

19.41 

18.92 

14.27 

19.64 

151 

6.97 

43.33 

12.94 

1.89 

20.09 

15.51 

15.82 

13.13 

16.02 

152 

9.37 

55.91 

12.54 

4.43 

27.54 

21.95 

22.58 

17.67 

23.10 

153 

7.34 

38.54 

11.55 

2.67 

16.82 

16.30 

17.98 

13.42 

16.67 

154 

7.27 

46.39 

15.76 

3.06 

20.12 

19.02 

20.24 

14.30 

21.33 

155 

10.44 

55.47 

21.28 

• 

22.73 

19.02 

22.15 

17.31 

21.39 

156 

• 

35.98 

12.40 

3.11 

13.34 

14.49 

15.08 

11.33 

15.46 

157 

6.17 

44.18 

13.20 

2.64 

18.33 

16.99 

17.60 

13.46 

18.17 

158 

6.48 

46.43 

15.72 

3.25 

20.65 

17.94 

19.96 

15.10 

20.10 

159 

5.60 

40.23 

16.01 

2.55 

13.83 

13.97 

14.75 

11.31 

14.60 

160 

9.55 

56.48 

18.23 

3.10 

24.12 

22.15 

21.57 

15.72 

21.36 

161 

6.48 

50.77 

15.13 

3.35 

18.89 

18.49 

20.19 

15.65 

20.04 

162 

8.17 

43.51 

16.26 

3.85 

16.77 

15.90 

17.55 

12.91 

17.58 

163 

8.06 

38.85 

13.66 

2.54 

15.94 

16.65 

17.61 

13.79 

17.05 

164 

6.83 

43.52 

13.15 

2.57 

17.19 

15.96 

15.81 

13.28 

16.61 

165 

8.93 

50.70 

20.59 

3.42 

21.87 

22.02 

21.47 

15.73 

21.02 

166 

11.42 

56.64 

23.64 

3.74 

24.99 

20.79 

20.37 

16.60 

21.98 

167 

4.91 

44.99 

12.52 

2.36 

19.15 

16.56 

17.28 

13.48 

17.56 

168 

6.38 

39.44 

15.48 

2.99 

15.60 

15.53 

17.03 

11.84 

17.61 

169 

6.10 

44.85 

9.34 

2.85 

18.23 

15.27 

16.80 

13.33 

15.97 

170 

8.38 

43.48 

9.72 

2.89 

19.01 

13.08 

12.59 

11.76 

15.79 

171 

7.34 

37.57 

9.24 

2.67 

17.53 

14.04 

15.51 

11.04 

14.87 

172 

10.82 

51.54 

16.20 

2.68 

20.97 

21.30 

21.96 

16.85 

23.98 

173 

7.09 

49.87 

18.40 

• 

16.09 

18.82 

20.48 

14.73 

20.33 

174 

7.44 

43.88 

16.48 

2.71 

15.55 

17.51 

16.97 

12.69 

17.15 

175 

6.36 

37.50 

14.68 

2.46 

11.92 

14.89 

14.69 

11.88 

14.49 

176 

9.91 

50.60 

9.04 

2.68 

25.05 

18.37 

20.86 

17.42 

21.57 

177 

7.14 

34.80 

11.86 

2.83 

14.53 

12.65 

13.27 

10.84 

12.72 

178 

8.54 

47.36 

15.12 

2.91 

18.07 

18.60 

18.61 

15.57 

19.03 

179 

7.89 

45.14 

16.18 

2.36 

17.47 

17.07 

17.41 

13.72 

17.85 

180 

7.37 

47.48 

14.19 

2.24 

20.23 

18.54 

18.87 

15.20 

19.08 

181 

7.21 

41.06 

14.30 

2.33 

16.82 

16.01 

17.12 

13.81 

17.76 

182 

7.59 

44.46 

13.72 

3.53 

17.09 

14.94 

16.36 

12.43 

15.33 

183 

8.64 

47.57 

16.07 

2.79 

19.16 

19.99 

19.67 

16.73 

19.92 

184 

8.13 

43.76 

14.18 

2.86 

17.68 

15.09 

15.89 

12.95 

16.10 

185 

8.86 

52.53 

10.74 

3.43 

21.66 

19.23 

19.71 

15.70 

19.95 

186 

6.58 

42.76 

10.83 

2.63 

17.19 

14.90 

16.14 

14.72 

17.24 

187 

8.33 

48.83 

15.51 

2.38 

17.78 

19.70 

20.53 

15.08 

21.02 

188 

9.09 

54.56 

19.49 

2.47 

19.65 

20.59 

20.35 

14.93 

20.58 

189 

6.18 

47.94 

14.13 

3.34 

19.72 

17.05 

18.33 

14.12 

18.11 

190 

8.30 

43.99 

16.52 

3.28 

17.29 

16.81 

18.93 

12.83 

18.47 

191 

6.03 

43.86 

13.93 

3.17 

16.64 

16.12 

16.82 

12.71 

16.59 

192 

6.26 

42.18 

14.96 

2.24 

17.23 

16.92 

17.37 

13.28 

18.07 


369 


# 

PPP 

POSAP 

ANLL 

ANLW 

ANLP 

PDI 

PDII 

PDIII 

PDIV 

193 

6.61 

47.70 

10.04 

2.80 

21.99 

19.92 

19.82 

15.54 

20.38 

194 

5.56 

33.08 

10.54 

2.29 

11.51 

12.43 

13.41 

10.65 

13.10 

195 

8.47 

45.28 

13.06 

2.99 

19.63 

20.02 

20.57 

16.24 

20.95 

196 

5.71 

38.76 

12.58 

2.47 

16.90 

15.59 

16.14 

12.33 

15.70 

197 

7.39 

48.36 

11.88 

2.67 

21.33 

19.48 

20.34 

15.98 

20,11 

198 

6.74 

41.10 

13.58 

3.03 

16.23 

18.56 

18.49 

12.83 

18.19 

199 

5.71 

42.81 

16.62 

2.46 

15.97 

17.53 

17.86 

13.23 

17.28 

200 

6.17 

42.95 

12.75 

2.19 

17.86 

17.42 

17.32 

14,68 

17.48 

201 

8.10 

43.93 

12.01 

3.20 

17.71 

17.32 

19.74 

14.12 

19.20 

202 

6.78 

46.30 

12.15 

3.06 

20.28 

19.92 

19.08 

13.72 

20.14 

203 

6.67 

35.02 

10.88 

2.74 

13,89 

12.81 

13.31 

11.07 

13.41 

204 

9.55 

54.78 

10.42 

3.21 

22.49 

22.77 

22.05 

14.75 

23.74 

205 

5.13 

32.88 

9.18 

2.92 

12.39 

11.23 

11.93 

10.26 

12.24 

206 

10.62 

59.79 

17.83 

4.44 

26.64 

21.48 

21.75 

17.34 

22.62 

207 

6.19 

42.50 

11.25 

2.71 

17.88 

14.51 

15.40 

11.98 

15.25 

208 

8.19 

41.54 

13.64 

2.79 

16.03 

14.73 

14.57 

11.43 

14.54 

209 

5.84 

29.71 

9.14 

2.57 

10.47 

11.21 

12.01 

9.82 

11.65 

210 

6.98 

33.05 

10.37 

2.11 

14.55 

12.21 

13.52 

10.40 

12.55 

211 

8.98 

42.77 

16.63 

2.61 

17.63 

18.79 

18.44 

14.78 

18.41 

212 

7.63 

44.20 

12.63 

3.20 

17.68 

17.93 

18.23 

13.55 

18.45 

213 

7.78 

45.79 

14.69 

2.69 

20.06 

16.68 

17.65 

14.34 

17.99 

214 

8.79 

49.21 

19.36 

2.17 

18.34 

19.87 

19.59 

15.14 

19.71 

215 

8.80 

45.96 

17.12 

2.38 

16.51 

17.60 

18.26 

14.92 

18.19 

216 

9.20 

47.16 

15.26 

2.72 

21,66 

19.30 

24.47 

14.92 

19.75 

217 

8.46 

46.80 

13.16 

2.67 

18.45 

17.31 

17.84 

14.31 

18.21 

218 

9.92 

49.01 

12.59 

3.11 

21.24 

18.16 

19.39 

15.58 

18.98 

219 

8.75 

47.92 

11.83 

3.44 

19.49 

18.63 

18.81 

13.14 

17.78 

220 

7.58 

41.93 

11.89 

2.47 

15.64 

14.91 

15.79 

12.64 

15.96 

221 

7.98 

42.26 

12.20 

2.81 

17.65 

15.80 

15.85 

12.41 

16,58 

222 

9.76 

50.92 

19.40 

2,62 

19.02 

20.53 

20.64 

15.28 

20.55 

223 

• 

45.12 

18.09 

3.68 

17.91 

18.69 

22.02 

14.03 

20.72 

224 

• 

52.98 

26.37 

3,53 

19.95 

23.72 

21.34 

16.80 

22.39 

225 

• 

48.42 

18.84 

3.81 

17.15 

20.04 

21.67 

13.85 

21.60 

226 

5.75 

30.75 

10.56 

2.83 

13.00 

11.11 

13.44 

9.23 

13.38 

227 

11.69 

• 

31.39 

3.51 

• 

24.49 

28.52 

20.67 

25.47 

228 

11.63 

58.91 

25.52 

3.44 

25.74 

24.33 

25.60 

17.84 

26.45 

229 

10.46 

54.21 

25.08 

3.45 

19.12 

21.09 

22.52 

16.35 

23.07 

230 

7.73 

45.79 

17.98 

3.78 

17.41 

17.78 

19.76 

12.91 

19.48 

231 

9.22 

53.72 

27.96 

3.21 

18.49 

20.24 

21.28 

15.29 

21.69 

232 

7.23 

42.10 

16.09 

• 

16.20 

15.92 

16.91 

13.14 

16.71 

233 

12.02 

39.03 

18.40 

8.39 

19.43 

12.81 

11.64 

8.28 

11.79 

234 

• 

40.52 

21.12 

7.46 

18.49 

13.63 

13.05 

9.49 

11.82 

235 

13.08 

39.53 

22.68 

7.41 

17.85 

12.85 

11.31 

8.55 

11.18 

236 

13.51 

38.07 

19.62 

7.62 

16.45 

12.89 

10.54 

8.34 

11.22 

237 

13.98 

35.58 

17.91 

5.20 

18.68 

11.72 

10.35 

8.43 

10.22 

238 

10.05 

28.01 

18.87 

7.55 

13.77 

9.60 

9.14 

6.99 

8.87 

239 

8.76 

25.10 

12.14 

6.55 

11.15 

8.20 

6.70 

5.47 

6.51 

240 

10.56 

31.57 

18.55 

8.16 

14,57 

10.10 

10.59 

8.67 

10.82 


370 


# 

PPP 

POSAP 

ANLL 

ANLW 

ANLP 

PDI 

PDII 

PDIII 

PDIV 

241 

9.74 

25.06 

13,13 

7.23 

12.05 

8.64 

8.12 

6.09 

7.96 

242 

9.01 

21.84 

13,83 

6.07 

10.40 

8.91 

7.72 

5.80 

7.41 

243 

• 

20.53 

9.43 

4.91 

10.87 

8.28 

7.18 

• 

7.81 

244 

6.73 

20.07 

7.72 

4.01 

8.90 

8.33 

6.17 

5.11 

6.40 

245 

7.16 

18.59 

8.68 

5.13 

9.23 

8,00 

6.51 

4.57 

6.23 

246 

6.23 

18.02 

9.22 

4.77 

7.72 

6.50 

6.12 

4.78 

6.13 

247 

6.61 

17,50 

7.45 

4.72 

8.48 

6.57 

5.76 

• 

6.31 

248 

6.57 

17.51 

7.72 

4.97 

8.86 

6.40 

5.85 

3.74 

5.88 

249 

6.09 

16.34 

9.85 

5.47 

7.70 

7.03 

5.88 

• 

5,99 

250 

6.22 

15.65 

8.59 

5.90 

7.62 

7.91 

5,57 

4.38 

5.75 

251 

6.75 

17.56 

7.34 

4.45 

8.06 

7.44 

6.45 

• 

5.82 

252 

6,18 

15.41 

6,55 

5.11 

7.79 

5.47 

4.80 

3.78 

5.35 

253 

6.10 

15.59 

7.49 

4.76 

7.17 

6.46 

5.33 

3.89 

5.49 

254 

4.88 

10.83 

3.37 

2.46 

6.17 

4.27 

4.00 

• 

4.00 

255 

6.43 

15.13 

7.89 

4.33 

7.18 

5.65 

4.71 

4.19 

4.97 

256 

5.80 

14.02 

5.58 

4.39 

6.88 

6.07 

5,33 

4.19 

4.72 

257 

5.88 

14.63 

5.46 

3.30 

7.37 

4.57 

4.39 

3.55 

4.88 

258 

5.79 

13.90 

8.15 

4.00 

6.61 

4.95 

4.55 

3.54 

4.44 

259 

5.44 

12.68 

5.30 

4.36 

6.60 

4.83 

3.98 

3.22 

3.81 

260 

5.13 

12.89 

4.90 

3.35 

6.70 

4.61 

4.29 

• 

4.22 

261 

5,01 

12.86 

5.86 

3.86 

5.81 

5.88 

4.40 

3.32 

4.55 

262 

4.14 

9.84 

3.31 

2.32 

4.41 

• 

• 

• 

• 

263 

10.22 

29.17 

16.28 

7.25 

12.77 

11.85 

9.65 

8.11 

9.81 

264 

12,64 

31.15 

18.14 

6.58 

14.85 

11.15 

9.85 

9.02 

9.65 

265 

9.23 

23,42 

12.47 

6.65 

10.88 

10.06 

8.68 

• 

7.38 

266 

• 

44.52 

20.53 

9.34 

21.57 

14.54 

12.74 

10.24 

12.62 

267 

14.34 

41.96 

24,99 

8.12 

18.56 

17.47 

13.44 

11.11 

13.33 

268 

17,59 

45.82 

25.89 

11.11 

24.23 

16,85 

14.39 

13.78 

14.99 

269 

17,40 

44.42 

20.31 

9.09 

22.32 

15,09 

12.77 

11.44 

13.14 

270 

16.90 

40,46 

18.66 

• 

24.39 

16,33 

11.98 

10.09 

11.45 

271 

• 

41.35 

22.39 

9.21 

16.63 

14.68 

12.50 

10.51 

11.84 

272 

11.72 

32.44 

21.32 

6.72 

14.67 

11.31 

9.93 

8.08 

10.01 

273 

10.05 

30.81 

17.46 

6.55 

12,91 

10.97 

8.99 

6.93 

9.38 

274 

9.59 

27.39 

12.52 

5.49 

12.52 

9.88 

8.22 

6.97 

8.82 

275 

9.72 

29.06 

13.40 

9.25 

12.98 

10.98 

9.52 

7.61 

9.82 

276 

7.20 

18.44 

9.83 

5.63 

8.78 

7.26 

6.53 

4.65 

6.53 

277 

11,02 

30.96 

13.72 

6.96 

13.98 

11.37 

8.77 

7.45 

• 

278 

9.67 

27.31 

11.70 

5.30 

12.76 

8.23 

7.36 

6.46 

7.76 

279 

7.72 

19.87 

12.72 

3.92 

10.06 

7.36 

6.92 

4.70 

6.58 

280 

10.55 

27.98 

16.42 

5.89 

12.87 

11.43 

8.65 

7.02 

8.89 

281 

8.51 

23.72 

11.64 

5.94 

10.16 

8.45 

7.53 

6.81 

7.92 

282 

9.34 

27.76 

16,67 

7.45 

13.05 

11.97 

9.55 

7.35 

9.22 

283 

9.30 

26.30 

15.82 

6.78 

13.81 

11,21 

9.01 

7,45 

8.08 

284 

9.32 

27.92 

13.59 

7.29 

11.62 

8.66 

7.84 

5.76 

7.16 

285 

8.42 

21,78 

9.02 

5.80 

9.81 

8.01 

6.61 

5.20 

6.48 

286 

9.35 

26.09 

13.68 

6,63 

12.12 

10.07 

8.13 

7.05 

8.08 

287 

• 

24.18 

14.18 

4.71 

10.80 

8.55 

7,52 

6.44 

7.70 

288 

8.06 

22.57 

9.29 

5.48 

10.09 

8.16 

7.18 

6.30 

7.18 


371 


# 

PPP 

POSAP 

ANLL 

ANLW 

ANLP 

PDI 

PDII 

PDIII 

PDIV 

289 

7.41 

20.40 

10.63 

5.01 

9.17 

7.81 

6.02 

5.20 

6.52 

290 

10.16 

24.13 

12.82 

7.81 

11.28 

9.63 

8.30 

6.32 

7.87 

291 

7.27 

20.35 

11.92 

5.62 

8.68 

7.77 

6.96 

4.98 

6.75 

292 

• 

18.89 

9.01 

5.35 

8.43 

6.81 

5.99 

• 

6.46 

293 

8.13 

18.89 

13.15 

6.89 

9.92 

7.76 

6.81 

4.66 

6.67 

294 

6.85 

18.28 

10.22 

6.03 

8.91 

8.19 

5.76 

4.76 

6.77 

295 

8.02 

22.62 

9.95 

6.31 

10.31 

9.19 

7.45 

6.13 

7.05 

296 

8.03 

20.60 

8.21 

5.02 

9.04 

7.01 

6.06 

5.12 

5.81 

297 

6.41 

18.36 

6.84 

4.74 

7.65 

6.11 

5.18 

• 

5.86 

298 

7.59 

19.33 

7.98 

5.89 

9.29 

8.06 

6.50 

4.66 

6.42 

299 

7.09 

19.18 

10.10 

5.44 

8.63 

7.57 

7.24 

6.03 

7.18 

300 

6.22 

16.95 

7.52 

4.74 

7.42 

6.72 

5.96 

5.17 

5.83 

301 

6.45 

16.84 

7.65 

4.60 

8.54 

7.13 

5.15 

4.62 

5.38 

302 

6.09 

15.79 

7.79 

4.54 

7.03 

6.28 

5.33 

4.02 

5.54 

303 

5.86 

16.59 

9.31 

5.09 

7.33 

6.47 

5.63 

4.10 

5.56 

304 

5.93 

14.17 

5.96 

3.65 

7.33 

6.33 

4.57 

3.77 

4.49 

305 

5.68 

15.15 

8.05 

4.67 

6.65 

6.43 

5.62 

3.98 

5.36 

306 

5.84 

14.95 

6.52 

4.35 

7.40 

5.15 

4.52 

3.18 

4.31 

307 

5.28 

15.56 

5.55 

3.03 

7.74 

5.96 

5.07 

• 

5.14 

308 

6.02 

15.01 

7.04 

4.01 

7.02 

5.96 

5.07 

4.29 

5.49 

309 

5.90 

17.08 

6.79 

5.27 

9.22 

7.07 

5.81 

4.32 

5.59 

310 

5.82 

16.05 

9.12 

5.15 

7.56 

7.52 

6.06 

4.69 

6.06 

311 

5.78 

15.29 

7.69 

5.14 

6.75 

5.64 

4.65 

3.63 

4.66 

312 

6.17 

16.38 

8.92 

4.76 

7.15 

7.64 

5.21 

4.24 

5.67 

313 

6.72 

17.99 

8.25 

4.40 

8.30 

5.86 

5.46 

4.90 

5.70 

314 

6.78 

19.35 

9.75 

5.14 

9.00 

7.47 

6.79 

4.84 

6.72 

315 

5.33 

13.86 

6.75 

3.78 

6.48 

6.64 

4.87 

3.77 

4.68 

316 

5.59 

14.14 

6.78 

4.72 

6.84 

4.81 

4.46 

3.50 

4.30 

317 

6.69 

16.64 

8.73 

5.42 

7.04 

6.74 

5.46 

5.45 

6.64 

318 

5.37 

14.29 

7.24 

3.98 

6.67 

5.95 

4.30 

2.84 

4.62 

319 

5.36 

13.14 

6.78 

4.20 

6.77 

5.45 

4.73 

3.87 

4.52 

320 

5.07 

13.64 

6.52 

4.29 

6.28 

4.85 

4.57 

3.18 

4.38 

321 

5.22 

12.69 

6.14 

3.30 

6.71 

5.25 

3.98 

• 

4.53 

322 

6.43 

15.86 

6.50 

4.35 

7.91 

5.40 

4.51 

3.91 

4.94 

323 

5.05 

12.03 

4.76 

2.97 

6.40 

5.17 

4.52 

• 

6.02 

324 

5.51 

12.95 

5.62 

3.47 

6.64 

5.50 

3.97 

3.44 

4.19 

325 

4.68 

11.36 

4.64 

3.24 

5.41 

4.05 

3.69 

2.70 

3.72 

326 

4.69 

10.22 

4.46 

3.46 

5.17 

4.26 

3.52 

• 

3.38 

327 

3.95 

10.68 

5.01 

3.20 

4.60 

4.40 

4.05 

2.96 

3.84 

328 

4.11 

10.21 

3.96 

2.78 

5.41 

4.06 

3.50 

2.60 

3.29 

329 

4.54 

10.03 

4.17 

2.54 

4.96 

3.32 

2.89 

• 

• 

330 

3.82 

9.03 

3.93 

2.97 

4.99 

3.33 

3.21 

• 

3.01 

331 

3.70 

9.78 

3.48 

2.46 

4.64 

3.55 

3.29 

• 

3.15 

332 

3.68 

9.25 

3.68 

2.62 

4.57 

• 

• 

• 

• 

333 

4.23 

8.93 

3.09 

2.28 

4.63 

• 

• 

• 

• 

334 

15.19 

41.58 

24.43 

7.17 

19.02 

14.99 

13.96 

9.22 

12.39 

335 

11.48 

30.26 

16.87 

8.39 

14.66 

10.10 

9.16 

6.30 

8.22 

336 

9.52 

24.43 

13.47 

7.09 

12.52 

11.47 

8.63 

6.67 

8.92 


372 


# 

PPP 

POSAP 

ANLL 

ANLW 

ANLP 

PDI 

PDII 

PDIII 

PDIV 

337 

8.86 

24.07 

13.55 

6.72 

10.76 

8.93 

7.78 

5.58 

8.03 

338 

7.90 

21.75 

12.36 

6.34 

9.62 

8.86 

7.56 

7.27 

8.02 

339 

8.94 

20.46 

12.29 

6.47 

9.70 

8.97 

7.29 

5.14 

6.90 

340 

8.13 

21.76 

9.56 

5.62 

9.86 

7.79 

6.82 

4.51 

6.78 

341 

7.39 

20.29 

9.52 

5.18 

9.13 

7.82 

6.99 

6.19 

6.44 

342 

5.80 

19.43 

9.95 

4.33 

8.43 

8.53 

6.72 

5.92 

6.14 

343 

6.37 

18.23 

9.16 

5.46 

9.32 

8.40 

6.48 

5.38 

6.43 

344 

7.43 

18.69 

8.96 

5.01 

8.17 

6.92 

6.52 

• 

5.46 

345 

6.76 

17.22 

7.54 

4.55 

8.42 

6.82 

5.86 

4.60 

5.95 

346 

6.40 

16.52 

7.08 

3.96 

7.78 

5.92 

5.50 

4.25 

4.98 

347 

5.89 

14.58 

7.29 

4.48 

6.56 

5.49 

4.78 

3.86 

4.97 

348 

5.09 

12.13 

4.94 

3.38 

5.95 

5.04 

4.65 

• 

4.53 

349 

4.69 

11.54 

4.51 

3.29 

5.79 

4.48 

3.29 

• 

3.44 

350 

4.71 

11.38 

3.17 

2.46 

6.15 

4.32 

3.43 

2.91 

3.37 

351 

4.19 

10.71 

3.97 

2.62 

5.33 

3.16 

3.55 

3.37 

3.82 

352 

4.16 

9.66 

3.29 

2.81 

4.63 

3.10 

2.77 

2.37 

3.22 

353 

3.64 

7.49 

2.72 

2.23 

3.92 

• 

• 

• 

• 

354 

7.63 

33.29 

7.97 

2.84 

13.89 

• 

• 

• 

12.77 

355 

• 



1.73 


• 

14.77 

12.77 

12.56 

356 

• 

• 


• 

• 

16.14 

14.43 

14.38 

• 

357 

• 

• 

18.50 

6.24 

• 

• 

• 

• 

• 

358 


• 

15.40 

7.45 

19.12 

• 

• 

• 

• 

359 


• 

12.12 

5.35 

• 

14.37 

16.49 

14.37 

16.35 

360 

• 

• 

12.96 

5.57 

• 

• 

• 

• 

• 

361 


41.74 

18.83 

6.64 

19.86 

• 

• 

• 

• 

362 

6.05 

32.41 

7.63 

3.37 

14.89 

12.14 

12.42 

10.17 

12.21 

363 

• 

29.93 

8.52 

3.86 

14.36 

10.33 

• 

• 

• 

364 

14.24 

30.75 

21.31 

11.89 

16.31 

• 

• 

• 

• 

365 

• 

26.35 

15.25 

11.13 

16.39 

7.02 

• 

• 

6.98 

366 

6.31 

11.65 

7.22 

5.99 

7.68 


• 

• 

• 

367 

5.47 

11.51 

6.56 

5.21 

6.21 

• 

• 

• 

• 

368 

• 

11.58 

6.55 

5.47 

5.68 


• 

• 

• 

369 

5.25 

10.92 

5.54 

5.52 

5.67 


• 

• 

• 

370 

13.43 

33.47 

22.77 

13.81 

21.46 

9.30 

9.28 

6.56 

9.25 

371 

• 

44.89 

12.24 

2.29 

20.30 

14.13 

13.98 

13.57 

16.97 

372 

• 

• 

32.94 

1.84 

• 

22.41 

21.83 

22.31 

25.90 


373 


# 

PDV 

TWMX 

LTH1 

LTH2 

LTH3 

LTH4 

LTH5 

TTH1 

TTH2 

1 

13.56 

76.27 

1.93 

6.33 

7.31 

5.75 

1.62 

1.87 

5.27 

2 

8.08 

45.87 

1.51 

3.41 

5.02 

3.53 

1.43 

1.46 

3.09 

3 

7.13 

34.41 

1.19 

2.76 

3.81 

2.75 

1.21 

1.04 

2.23 

4 

6.01 

32.82 

1.49 

2.71 

3.45 

2.52 

1.13 

1.06 

2.19 

5 

5.52 

27.81 

1.02 

2.24 

3.23 

1.70 

0.90 

1.00 

1.84 

6 

• 

21.97 

1.04 

1.86 

2.46 

1.68 

0.87 

0.87 

1.53 

7 

12.59 

73.27 

1.79 

5.58 

6.80 

4.99 

1.54 

1.65 

4.87 

8 

7.17 

45.65 

1.27 

3.18 

4.49 

2.92 

1.24 

1.16 

2.80 

9 

6.38 

43.05 

1.27 

3.30 

3.91 

3.11 

1.27 

1.20 

2.69 

10 

8.39 

45.46 

1.85 

3.75 

5.15 

3.12 

1.60 

1.42 

3.08 

11 

6.57 

40.28 

1.24 

3.08 

4.52 

2.81 

1.16 

1.23 

2.46 

12 

6.75 

35.10 

1.24 

2.87 

3.86 

2.83 

0.96 

1.02 

2.54 

13 

6.55 

31.39 

1.05 

2.75 

3.65 

2.13 

1.23 

1.19 

2.23 

14 

5.58 

28.90 

0.63 

2.15 

3.27 

2.13 

0.81 

0.99 

1.67 

15 

• 

21.45 

0.91 

1.81 

• 

1.85 

0.91 

0.91 

1.66 

16 

• 

• 

1.34 

2.33 

3.50 

2.87 

1.15 

• 

2.23 

17 

13.06 

73.92 

2.29 

5.86 

7.82 

7.04 

1.73 

1.87 

5.47 

18 

10.71 

64.73 

1.60 

4.77 

6.31 

5.00 

1.48 

1.47 

4.21 

19 

11.25 

62.92 

1.72 

4.35 

6.23 

4.72 

1.53 

1.42 

3.45 

20 

13.53 

76.07 

2.08 

6.24 

8.03 

6.19 

1.77 

1.66 

5.18 

21 

6.29 

38.15 

0.83 

2.76 

3.81 

2.45 

1.09 

1.05 

1.98 

22 

5.29 

27.78 

1.11 

2.36 

3.51 

2.15 

1.13 

1.11 

1.90 

23 

6.62 

35.15 

1.46 

2.92 

3.88 

2.59 

1.25 

1.38 

2.41 

24 

6.73 

34.59 

1.14 

2.83 

3.82 

2.33 

1.04 

0.88 

2.19 

25 

6.37 

• 

1.07 

2.95 

3.72 

2.95 

1.02 

1.05 

2.42 

26 

• 

• 

1.04 

2.27 

3.83 

2.22 

0.96 

• 

2.09 

27 

6.35 

33.87 

0.80 

2.51 

3.32 

2.24 

• 

0.95 

2.00 

28 

10.51 

57.41 

1.54 

3.75 

5.94 

4.61 

1.51 

1.27 

3.39 

29 

7.34 

40.65 

1.37 

3.22 

4.59 

2.98 

1.19 

1.25 

2.65 

30 

• 

• 

0.62 

2.47 

3.30 

2.29 

1.13 

1.23 

2.15 

31 

• 

29.69 

0.80 

1.91 

2.76 

• 

• 

0.96 

1.72 

32 

4.99 

24.19 

0.96 

2.03 

2.81 

1.63 

0.95 

0.86 

1.63 

33 

4.01 

20.21 

0.63 

1.46 

2.08 

1.49 

0.87 

0.81 

1.24 

34 

4.03 

19.30 

0.81 

1.75 

2.23 

1.46 

0.95 

0.92 

1.40 

35 

10.87 

64.02 

1.67 

4.96 

6.79 

5.48 

1.68 

1.39 

4.31 

36 

10.23 

58.72 

1.52 

4.40 

5.70 

4.83 

0.99 

1.32 

3.73 

37 

11.15 

66.54 

1.66 

5.14 

6.52 

5.35 

1.09 

1.57 

4.33 

38 

6.43 

34.93 

0.73 

2.47 

3.32 

2.37 

0.86 

1.02 

2.17 

39 

6.24 

33.12 

1.25 

2.61 

3.86 

2.45 

1.11 

1.18 

2.09 

40 

• 

30.45 

0.88 

2.61 

3.59 

2.59 

1.15 

0.97 

2.31 

41 

5.23 

30.79 

0.74 

2.19 

2.99 

2.12 

0.87 

0.95 

1.86 

42 

5.38 

29.48 

1.19 

2.46 

• 

2.43 

0.92 

1.10 

2.12 

43 

6.76 

28.72 

0.96 

2.26 

3.50 

1.96 

0.71 

0.85 

1.68 

44 

5.46 

27.15 

1.33 

2.13 

3.23 

2.14 

0.96 

0.99 

1.77 

45 

4.48 

21.75 

0.82 

2.19 

2.93 

1.91 

0.93 

0.97 

1.72 

46 

12.59 

77.29 

2.22 

5.44 

• 

5.77 

1.43 

1.63 

4.83 

47 

5.81 

31.19 

0.93 

2.37 

3.60 

2.38 

1.02 

1.04 

1.90 

48 

5.16 

25.55 

1.01 

2.14 

3.06 

1.84 

1.02 

0.88 

1.76 


374 


# 

PDV 

TWMX 

LTH1 

LTH2 

LTH3 

LTH4 

LTH5 

TTH1 

TTH2 

49 

4.74 

26.49 

0.69 

1.71 

2.38 

1.62 

0.74 

0.82 

1.38 

50 

4.77 

26.21 

0.88 

2.04 

2.75 

2.03 

0.95 

0.95 

1.71 

51 

3.93 

22.96 

0.83 

1.79 

2.37 

1.90 

0.72 

0.77 

1.75 

52 

• 

23.92 

0.66 

2.05 

2.51 

1.87 

1.01 

0.85 

1.82 

53 

• 

19.43 

0.97 

1.75 

2.43 

1.85 

0.85 

0.85 

1.60 

54 

• 

• 

0.71 

1.77 

2.24 

1.81 

0.83 

0.86 

1.46 

55 

• 

18.77 

0.77 

1.56 

2.09 

1.43 

0.72 

0.64 

1.32 

56 

• 

17.65 

0.66 

1.53 

2.10 

1.48 

0.67 

0.64 

1.15 

57 

• 

14.85 

0.64 

0.96 

1.67 

1.10 

0.69 

0.72 

0.92 

58 

8.63 

42.51 

1.46 

3.25 

5.02 

3.43 

1.46 

1.46 

2.86 

59 

9.16 

42.51 

1.35 

2.76 

4.23 

3.03 

1.21 

1.34 

2.15 

60 

• 

58.36 

1.74 

3.98 

6.18 

4.07 

1.81 

1.71 

3.48 

61 

7.27 

39.24 

1.55 

3.32 

4.54 

3.25 

1.32 

1.27 

2.65 

62 

6.21 

33.15 

0.73 

2.82 

• 

2.53 

1.33 

1.12 

2.17 

63 

7.63 

45.46 

1.30 

3.38 

4.63 

3.40 

1.37 

1.35 

2.71 

64 

7.08 

33.27 

1.46 

2.68 

4.05 

2.83 

1.25 

1.32 

2.23 

65 

6.90 

34.79 

1.07 

2.54 

3.89 

2.61 

1.25 

1.18 

2.07 

66 

13.67 

77.06 

2.29 

5.95 

7.09 

5.79 

1.85 

2.01 

5.15 

67 

7.67 

35.54 

0.99 

2.79 

3.63 

2.92 

1.09 

1.15 

2.58 

68 

7.67 

42.62 

1.73 

3.70 

• 

3.72 

1.58 

1.50 

2.90 

69 

10.07 

54.85 

1.56 

4.83 

5.94 

4.06 

1.54 

1.59 

3.57 

70 


34.56 

1.27 

3.40 

3.80 

2.79 

1.15 

1.30 

2.74 

71 

7.18 

39.95 

1.49 

3.41 

4.77 

3.34 

1.57 

1.34 

2.63 

72 

6.88 

33.61 

1.17 

2.58 

3.23 

2.19 

1.26 

1.19 

2.07 

73 


62.81 

1.89 

• 

• 

• 

• 

1.65 

4.23 

74 

7.28 

37.94 

1.29 

3.17 

4.11 

2.93 

1.28 

1.44 

2.52 

75 

5.81 

37.99 

0.98 

2.54 

3.11 

2.54 

1.09 

1.19 

2.35 

76 

7.41 

42.08 

1.67 

3.78 

4.60 

3.59 

1.63 

• 

• 

77 

13.91 

80.51 

2.25 

5.61 

• 

• 

1.63 

1.84 

5.47 

78 

• 

28.34 

0.85 

2.18 

2.89 

1.89 

0.95 

0.91 

1.70 

79 

5.95 

31.10 

0.96 

2.52 

3.52 

2.54 

1.30 

1.24 

2.10 

80 

5.48 

28.48 

0.70 

2.74 

3.26 

2.05 

1.01 

1.22 

2.08 

81 

4.76 

26.68 

1.15 

2.01 

2.58 

1.85 

1.03 

1.01 

1.55 

82 

4.96 

28.66 

1.03 

2.26 

3.14 

2.14 

1.20 

0.98 

1.88 

83 

5.32 

25.13 

0.76 

1.87 

• 

1.69 

1.11 

0.90 

1.49 

84 

5.72 

29.37 

1.09 

2.59 

3.52 

2.36 

1.22 

1.25 

2.07 

85 

6.78 

• 

1.24 

2.44 

3.99 

2.71 

1.09 

• 

• 

86 

6.50 

• 

1.44 

2.89 

3.54 

2.72 

1.23 

• 

• 

87 

6.22 

31.90 

1.12 

2.52 

3.78 

2.58 

1.23 

1.18 

2.06 

88 

7.08 

42.41 

1.08 

3.66 

4.73 

3.37 

1.29 

1.56 

2.96 

89 

13.57 

79.37 

2.36 

5.74 

7.85 

6.59 

1.57 

1.86 

5.04 

90 

5.39 

26.74 

0.73 

2.49 

3.27 

2.19 

1.01 

• 

• 

91 

5.13 

29.69 

1.27 

2.49 

3.28 

2.20 

1.08 

1.22 

1.66 

92 

5.25 

28.21 

0.84 

1.97 

2.91 

2.10 

1.06 

1.17 

1.75 

93 

4.44 

27.21 

0.97 

1.88 

2.35 

1.58 

0.82 

0.93 

1.42 

94 

5.15 

27.33 

0.81 

2.10 

2.97 

2.12 

1.22 

1.08 

1.60 

95 

5.28 

26.82 

1.08 

2.51 

3.10 

2.12 

1.12 

1.06 

1.99 

96 

5.48 

28.18 

0.78 

2.18 

2.98 

2.25 

1.25 

1.17 

1.76 


375 


# 

PDV 

TWMX 

LTH1 

LTH2 

LTH3 

LTH4 

LTH5 

TTH1 

TTH2 

97 

4.74 

26.60 

1.09 

2.18 

2.92 

2.06 

1.10 

1.07 

1.74 

98 

14.24 

79.51 

2.49 

6.39 

8.30 

7.33 

1.83 

1.87 

5.56 

99 

10.74 

60.12 

1.99 

4.42 

6.26 

5.23 

1.45 

1.61 

3.96 

100 

12.20 

71.75 

2.10 

5.23 

7.18 

5.82 

1.64 

1.68 

4.77 

101 

10.19 

55.95 

1.68 

3.77 

• 

4.16 

1.49 

1.50 

3.47 

102 

10.78 

63.46 

1.91 

3.93 

5.92 

4.59 

1.32 

1.35 

3.33 

103 

5.71 

30.78 

1.11 

2.53 

3.41 

2.58 

1.13 

1.10 

1.99 

104 

• 

26.47 

0.97 

2.09 

2.58 

2.12 

0.94 

0.91 

1.71 

105 

6.10 

27.59 

0.79 

2.67 

3.43 

2.25 

1.07 

0.92 

1.98 

106 

4.83 

27.62 

1.09 

2.28 

3.07 

2.07 

1.25 

1.12 

1.74 

107 

5.79 

30.27 

0.88 

2.36 

3.28 

2.36 

1.24 

1.15 

1.99 

108 

4.91 

27.77 

1.19 

2.51 

3.05 

2.04 

1.43 

1.17 

2.16 

109 

4.45 

25.97 

0.94 

2.24 

2.75 

2.09 

0.94 

1.01 

1.71 

110 

5.74 

31.05 

1.26 

2.48 

3.63 

2.36 

1.25 

1.33 

2.14 

111 

5.11 

27.91 

0.89 

2.49 

3.06 

2.12 

1.25 

1.20 

1.97 

112 

4.27 

22.39 

0.76 

1.76 

2.63 

1.59 

0.96 

0.95 

1.46 

113 

3.68 

23.31 

1.08 

1.90 

2.52 

1.99 

0.89 

0.90 

1.71 

114 

5.38 

29.85 

1.26 

2.56 

3.37 

2.26 

1.25 

1.27 

2.09 

115 

« 

21.76 

0.94 

2.04 

2.46 

1.93 

1.06 

0.93 

1.67 

116 

4.82 

28.98 

1.09 

2.23 

3.22 

2.35 

1.24 

1.08 

1.78 

117 

• 

20.85 

0.99 

1.73 

2.30 

1.50 

0.85 

0.91 

1.29 

118 

3.36 

18.55 

1.05 

1.51 

2.13 

1.58 

0.98 

0.85 

1.42 

119 

3.14 

18.14 

0.88 

1.84 

2.18 

1.67 

0.91 

0.88 

1.49 

120 

11.82 

66.57 

1.95 

4.83 

6.98 

5.35 

1.60 

1.80 

4.47 

121 

8.31 

49.03 

1.72 

3.82 

5.15 

4.06 

1.20 

1.42 

3.14 

122 

5.16 

28.35 

« 

2.40 

3.06 

2.36 

1.10 

1.22 

2.17 

123 

• 

25.51 

0.80 

2.06 

2.52 

2.05 

1.06 

1.08 

1.76 

124 

• 

22.97 

1.01 

2.26 

2.76 

2.12 

1.30 

1.10 

1.84 

125 

• 

23.32 

0.79 

2.07 

2.70 

1.99 

1.07 

0.95 

1.67 

126 

3.83 

19.49 

1.02 

1.73 

2.28 

1.65 

0.93 

0.90 

1.50 

127 

14.21 

84.84 

2.31 

5.83 

• 

6.95 

1.64 

1.78 

4.97 

128 

• 

38.15 

0.73 

3.45 

3.98 

3.23 

1.26 

1.26 

2.99 

129 

11.64 

77.80 

2.37 

6.13 

7.78 

6.37 

1.67 

1.88 

4.97 

130 

• 

105.95 

2.60 

7.91 

• 

8.08 

• 

2.28 

6.27 

131 

• 

60.32 

2.17 

6.57 

7.89 

5.74 

1.06 

1.70 

6.08 

132 

11.02 

70.48 

3.84 

8.22 

9.08 

6.97 

2.08 

3.18 

7.56 

133 

10.21 

• 

1.82 

7.16 

8.45 

6.15 

2.08 

2.61 

6.09 

134 

12.83 

• 

1.58 

3.96 

5.19 

2.99 

• 

1.11 

3.96 

135 

10.97 

59.24 

3.16 

6.17 

8.20 

6.29 

2.12 

2.43 

5.28 

136 

9.98 

• 

2.33 

5.80 

• 

5.48 

1.56 

2.04 

5.43 

137 

3.68 

22.96 

0.83 

2.04 

2.56 

1.89 

1.04 

0.99 

1.72 

138 

10.49 

69.29 

3.12 

6.36 

8.17 

6.32 

1.87 

2.29 

5.48 

139 

9.27 

64.69 

3.02 

5.66 

7.40 

5.47 

1.43 

1.79 

5.29 

140 

9.51 

56.10 

2.59 

5.84 

7.08 

5.46 

1.71 

1.90 

5.14 

141 

• 

21.18 

1.47 

2.04 

2.76 

1.94 

0.99 

1.14 

2.13 

142 

10.87 

65.68 

3.32 

6.87 

8.48 

6.05 

2.20 

2.59 

6.07 

143 

16.17 

129.33 

3.04 

• 

• 

8.86 

1.89 

2.29 

7.02 

144 

• 

25.71 

1.02 

2.05 

2.64 

2.10 

1.10 

1.01 

1.79 


376 


# 

PDV 

TWMX 

LTH1 

LTH2 

LTH3 

LTH4 

LTH5 

TTH1 

TTH2 

145 

5.54 

31.71 

1.02 

2.94 

3.34 

2.69 

1.14 

1.18 

2.41 

146 

• 

30.39 

1.47 

• 

• 

2.60 

1.04 

1.27 

• 

147 

12.14 

73.10 

2.97 

8.39 

10.56 

8.16 

2.56 

3.25 

8.00 

148 

• 

65.60 

2.87 

6.42 

8.35 

6.36 

1.95 

2.23 

6.27 

149 

18.07 

114.60 

1.18 

8.48 

• 

8.02 

1.28 

1.53 

5.82 

150 

19.73 

104.03 

1.82 

6.99 

• 

5.18 

0.92 

1.29 

5.20 

151 

15.59 

98.62 

1.79 

6.79 

• 

5.86 

1.09 

1.38 

4.25 

152 

21.68 

122.01 

2.59 

8.50 

• 

9.54 

1.24 

1.61 

6.30 

153 

16.28 

85.21 

1.83 

6.32 

• 

5.32 

1.01 

1.33 

4.10 

154 

18.88 

104.59 

2.00 

8.01 

• 

6.50 

1.23 

1.38 

5.49 

155 

19.40 

120.76 

2.01 

10.50 

• 

9.07 

1.50 

1.64 

7.30 

156 

13.84 

76.39 

1.57 

5.55 

7.77 

4.98 

1.04 

• 

3.82 

157 

17.17 

• 

1.47 

6.08 

9.01 

5.20 

1.21 

1.18 

4.31 

158 

18.08 

107.46 

1.54 

7.60 

• 

5.72 

0.96 

1.24 

5.14 

159 

13.91 

89.78 

1.32 

5.32 

• 

5.35 

0.90 

1.21 

4.24 

160 

21.55 

120.27 

2.52 

9.09 

• 

9.33 

1.26 

1.56 

6.43 

161 

19.32 

113.15 

1.57 

7.08 

• 

6.38 

1.06 

1.16 

5.33 

162 

16.76 

96.58 

1.94 

6.26 

• 

6.06 

1.15 

1.54 

4.67 

163 

16.02 

87.30 

1.80 

6.22 

• 

5.21 

1.12 

1.15 

4.22 

164 

16.11 

93.85 

1.10 

5.61 

• 

6.21 

1.10 

1.23 

4.26 

165 

21.32 

116.36 

2.70 

10.22 

• 

8.59 

1.22 

1.65 

6.99 

166 

20.08 


2.28 

10.45 

• 

8.90 

1.46 

• 

• 

167 

16.24 

96.93 

1.27 

6.78 

• 

5.99 

0.88 

1.14 

4.29 

168 

16.73 


1.47 

6.74 

9.55 

4.97 

0.96 

0.91 

4.81 

169 

16.40 

91.70 

1.32 

5.25 


4.57 

1.01 

1.06 

3.73 

170 

15.10 


1.65 

6.23 

• 

• 

• 

1.31 

5.61 

171 

13.87 

84.39 

1.59 

5.82 

8.66 

5.71 

1.18 

1.28 

4.09 

172 

21.57 

119.74 

2.38 

8.42 

• 

7.57 

1.27 

1.72 

6.08 

173 

19.38 

111.84 

1.98 

8.48 

• 

6.35 

1.27 

1.53 

6.15 

174 

17.45 

93.93 

1.65 

• 

• 

• 

• 

1.08 

5.23 

175 

14.64 

79.69 

1.27 

5.09 

8.58 

4.38 

0.96 

1.05 

3.58 

176 

19.39 

114.95 

1.84 

7.83 

• 

8.31 

1.39 

1.44 

6.91 

177 

13.07 

76.64 

1.56 

5.94 

8.96 

5.55 

1.18 

1.19 

4.44 

178 

17.34 

• 

• 

6.98 

10.36 

6.71 

0.96 

• 

• 

179 

17.30 

102.70 

1.73 

7.57 


7.37 

0.97 

1.20 

5.61 

180 

19.11 

108.39 

1.63 

7.03 


5.41 

0.93 

1.23 

4.78 

181 

17.34 

87.34 

1.58 

6.02 

9.40 

6.28 

1.04 

• 

• 

182 

16.21 

95.72 

1.54 

5.95 

• 

5.75 

1.13 

1.47 

4.86 

183 

19.89 

104.85 

1.63 

6.67 

• 

6.24 

1.05 

1.31 

5.07 

184 

15.80 

94.16 

1.95 

6.01 

• 

6.81 

1.06 

1.23 

4.52 

185 

18.79 

116.52 

2.13 

8.79 

• 

7.86 

1.16 

1.48 

6.72 

186 

15.55 

88.77 

1.84 

6.18 

• 

4.87 

0.98 

1.30 

3.89 

187 

20.07 

104.53 

2.19 

7.97 

• 

7.74 

1.08 

1.44 

5.91 

188 

20.17 

116.69 

2.59 

9.42 

• 

8.87 

1.31 

1.64 

6.80 

189 

17.74 

104.91 

1.60 

6.84 


6.69 

1.09 

1.35 

4.96 

190 

16.75 

102.06 

2.02 

7.38 

• 

7.05 

1.19 

1.46 

5.30 

191 

16.11 

98.66 

1.71 

6.19 

• 

5.43 

0.95 

1.04 

4.45 

192 

16.67 

91.46 

1.46 

5.66 

• 

6.19 

0.97 

1.09 

3.86 


377 


# 

PDV 

TWMX 

LTH1 

LTH2 

LTH3 

LTH4 

LTH5 

TTH1 

TTH2 

193 

19.77 

105,46 

1.81 

7.12 

• 

6.85 

0.87 

1.40 

5.28 

194 

12,45 

69.88 

1.11 

4.16 

7.23 

4,57 

0.78 

0.96 

3.55 

195 

20.03 

102.17 

2.18 

7.50 

• 

6,10 

1.09 

1.27 

5.37 

196 

15.34 

85.05 

1.63 

6.18 

• 

5.54 

0.90 

1,01 

4.02 

197 

19.36 

107.28 

1.27 

7.01 

• 

5.88 

1.11 

1.19 

5.08 

198 

18.02 

95.96 

1.43 

5.79 

• 

5.37 

0.99 

1.11 

4.24 

199 

17.09 

97.14 

1.68 

7.72 

• 

5.58 

1.00 

1.30 

5.08 

200 

16.85 

98,48 

1.57 

6.60 

• 

6.10 

1.01 

1.10 

4.53 

201 

17.38 

95.91 

2.01 

6.23 

• 

5,89 

1.16 

1.38 

4.51 

202 

19,74 

105.86 

1.67 

6,47 

• 

6.70 

1.01 

1.29 

4,78 

203 

12,52 

74.94 

1.80 

5.38 

7.54 

4.72 

1.10 

1.26 

3.95 

204 

23.27 

125.67 

1,93 

8.01 

• 

6.84 

1.21 

1.32 

5,72 

205 

11.74 

69.55 

1.09 

4.12 

6,89 

3.97 

0.90 

1.23 

3.21 

206 

22.42 

135.33 

2.94 

10.69 

• 

9.56 

1.44 

1.78 

8.55 

207 

14.45 

92.84 

1.28 

6,12 

• 

4.80 

0.96 

0.96 

3.87 

208 

15.64 

91.01 

1.86 

7.04 

8.97 

5.26 

1.24 

1.42 

4,50 

209 

11.26 

• 

1.29 

3.95 

5.88 

3.58 

0.88 

0.93 

2.49 

210 

12.18 

74.21 

1.29 

5.08 

7.67 

4.80 

0.88 

1.09 

3,72 

211 

17.66 

98.35 

1.84 

6.15 

« 

5.67 

0.97 

1.21 

4.05 

212 

17.82 

93.73 

1.73 

6.42 

• 

6.14 

0.98 

1.22 

4.97 

213 

16.47 

105.82 

1.92 

7.49 

• 

6.17 

1.10 

1,31 

5.55 

214 

19.47 

105.02 

2.42 

7,73 

• 

7.27 

1,18 

1.56 

5.98 

215 

17.25 

97.74 

1.85 

7.83 


6.86 

1.02 

1.18 

5.78 

216 

18.64 

107.14 

1.97 

• 


• 

• 

1.35 

4.99 

217 

17.73 

105.70 

1.63 

7.55 


5.73 

0.99 

1.37 

4.79 

218 

20.39 

107.91 

1.74 

7.85 

• 

8.42 

1.30 

1.38 

5.86 

219 

17.25 

104.02 

1.88 

8.46 


7.65 

1.25 

1.35 

6.35 

220 

15.14 

92.31 

1.51 

6.35 


5.27 

1.09 

1.33 

4.61 

221 

15.60 

94.96 

1.60 

6.73 


6.22 

0.98 

1.17 

4.83 

222 

20.53 

118.24 

1.93 

9,15 


7.34 

1.05 

1.53 

6.44 

223 

19.34 

106.73 

2.73 

10.80 


7.63 

1.21 

1.51 

6.73 

224 

24.33 

137.94 

2.31 

11.37 

• 

8.55 

1.53 

1.87 

7.12 

225 

21.92 

109.93 

2.84 

10.31 

• 

7.87 

1.51 

1.61 

7.48 

226 

11.96 

72.36 

2.12 

7.82 

• 

6.32 

1.30 

1.48 

5.85 

227 

24.84 

• 

2.85 

11.82 

• 

10,21 

2.04 

2.31 

8.14 

228 

26.73 

140.53 

• 

11.04 

• 

9.39 

1.79 

2.01 

8.63 

229 

21.65 

129.38 

2.89 

10.66 

• 

8.63 

1.86 

1.94 

8.00 

230 

18.05 

107.89 

2.08 

10.64 

• 

6.87 

1.10 

1.54 

6.90 

231 

21.46 

132.63 

2.52 

11.58 

• 

7.51 

1.47 

2.15 

6.78 

232 

15.97 

96.38 

2.23 

8.53 

• 

6.50 

1.16 

1.42 

6.40 

233 

13.39 

86.85 

3.23 

6,31 

8.44 

6.73 

3.06 

3.16 

5.57 

234 

13.30 

93.89 

4.06 

7.18 

• 

6.71 

2.47 

3.00 

6.59 

235 

13.41 

88.16 

3.72 

6.31 

9.20 

6.59 

2.99 

3.03 

6.08 

236 

12.54 

83.09 

3.94 

8.11 

10.73 

7.59 

3.83 

3.39 

7.27 

237 

11.35 

79.64 

4.02 

5.81 

7,77 

6.69 

3.03 

2.76 

5.44 

238 

10.38 

66.99 

3.59 

6.23 

7.98 

6.93 

2.62 

2.71 

5.66 

239 

8.63 

56.64 

3.54 

5.70 

7.63 

6.55 

3.12 

3.23 

5.67 

240 

10.92 

72.54 

3.92 

5.48 

7.62 

7.16 

2.94 

3.20 

5.61 


378 


# 

PDV 

TWMX 

LTH1 

LTH2 

LTH3 

LTH4 

LTH5 

TTH1 

TTH2 

241 

8.68 

57.41 

3.03 

4.92 

6.89 

5.96 

2.15 

2.59 

4.57 

242 

8.86 

49.53 

2.81 

4.39 

5.81 

4.82 

2.51 

2.42 

4.12 

243 

7.74 

46.22 

2.48 

4.24 

5.86 

5.46 

2.26 

1.99 

4.12 

244 

8.08 

42.87 

2.90 

4.35 

6.13 

5.20 

2.19 

2.08 

4.20 

245 

7.35 

40.94 

2.69 

3.86 

5.89 

5.28 

1.84 

2.15 

4.29 

246 

6.75 

40.38 

2.07 

3.79 

4.95 

4.25 

1.61 

1.79 

3.46 

247 

7.22 

38.32 

3.02 

4.29 

5.48 

4.72 

2.10 

2.38 

4.27 

248 

6.64 

37.38 

2.71 

4.02 

5.34 

4.83 

1.91 

2.36 

4.08 

249 

7.54 

36.91 

1.98 

3.51 

4.74 

3.60 

1.70 

1.56 

3.37 

250 

6.94 

37.90 

2.28 

3.49 

4.44 

3.81 

1.85 

1.99 

3.20 

251 

7.20 

36.52 

2.04 

3.91 

5.28 

4.57 

1.93 

1.93 

3.79 

252 

5.57 

33.99 

2.05 

3.65 

4.72 

3.64 

1.85 

2.05 

3.40 

253 

6.18 

34.74 

2.05 

3.09 

4.19 

3.65 

1.72 

1.66 

3.04 

254 

3.93 

25.05 

1.63 

2.59 

3.32 

2.51 

1.20 

1.46 

2.13 

255 

5.68 

34.50 

2.38 

3.59 

4.74 

3.82 

1.90 

1.82 

3.65 

256 

6.23 

31.11 

2.22 

3.12 

4.21 

3.14 

1.57 

2.03 

3.17 

257 

5.23 

31.10 

2.13 

3.42 

4.26 

3.58 

1.65 

1.95 

3.27 

258 

5.52 

29.99 

1.98 

3.25 

3.98 

3.20 

1.62 

1.95 

3.06 

259 

5.10 

29.01 

1.79 

2.76 

3.81 

3.09 

1.66 

1.86 

2.61 

260 

5.05 

28.53 

1.72 

2.60 

3.34 

2.75 

1.86 

1.54 

2.38 

261 

5.29 

27.20 

1.57 

2.59 

3.59 

2.67 

1.35 

1.32 

2.42 

262 

• 

20.82 

1.19 

2.15 

2.88 

2.37 

1.40 

1.43 

2.09 

263 

13.74 

68.39 

4.69 

6.45 

9.97 

7.74 

3.71 

3.89 

6.22 

264 

10.85 

70.46 

5.11 

6.75 

9.08 

8.93 

3.46 

3.89 

6.56 

265 

8.92 

52.42 

4.15 

5.06 

6.41 

5.79 

3.16 

3.29 

5.29 

266 

14.51 

99.50 

5.76 

8.87 


11.45 

3.43 

3.86 

9.42 

267 

16.79 

96.63 

5.57 

7.87 

• 

11.83 

4.11 

4.37 

8.09 

268 

16.72 

107.51 

4.92 

8.55 

• 

12.47 

4.03 

4.04 

8.11 

269 

15.86 

101.05 

4.59 

7.75 


9.91 

3.64 

4.01 

7.19 

270 

16.43 

99.83 

5.49 

8.83 

• 

10.54 

4.12 

• 

• 

271 

17.03 

94.32 

5.64 

7.62 


10.33 

3.61 

4.23 

7.88 

272 

11.65 

71.78 

4.43 

6.94 

9.39 

8.55 

4.43 

3.43 

6.91 

273 

12.08 

64.71 

4.53 

6.09 

7.41 

6.65 

2.85 

3.74 

5.83 

274 

9.91 

62.10 

4.78 

5.88 

7.37 

9.61 

4.19 

3.96 

6.37 

275 

12.35 

65.07 

5.14 

6.69 

9.07 

9.47 

3.69 

3.96 

6.61 

276 

7.03 

42.41 

3.51 

4.33 

5.66 

5.44 

2.55 

2.62 

4.07 

277 

11.17 

67.68 

5.08 

6.73 

8.57 

9.55 

3.59 

3.84 

6.23 

278 

9.17 

60.73 

4.44 

6.34 

8.13 

8.13 

3.59 

3.47 

6.01 

279 

7.61 

43.66 

3.75 

4.70 

5.83 

6.14 

2.93 

3.05 

4.69 

280 

11.15 

65.41 

3.90 

5.85 

7.93 

7.46 

3.08 

3.04 

5.18 

281 

8.64 

54.18 

3.74 

4.84 

6.17 

5.25 

2.98 

2.89 

4.64 

282 

11.08 

66.38 

4.35 

5.63 

7.83 

7.18 

2.75 

3.26 

5.56 

283 

11.51 

65.25 

4.02 

5.48 

7.56 

6.98 

2.91 

3.03 

4.83 

284 

8.81 

61.01 

4.48 

5.63 

7.51 

7.15 

3.83 

3.38 

5.34 

285 

8.72 

45.35 

3.78 

4.80 

6.12 

5.62 

2.75 

3.13 

4.84 

286 

10.85 

64.12 

3.69 

5.19 

6.99 

6.91 

2.76 

3.14 

4.62 

287 

8.33 

51.38 

3.75 

5.42 

6.94 

6.89 

2.76 

2.99 

4.79 

288 

8.66 

48.16 

3.74 

4.84 

6.19 

5.78 

2.71 

2.84 

4.69 


379 


# 

PDV 

TWMX 

LTH1 

LTH2 

LTH3 

LTH4 

LTH5 

TTH1 

TTH2 

289 

8.23 

46.21 

3.76 

4.56 

5.63 

5.08 

2.48 

2.68 

3.98 

290 

9.16 

57.69 

4.31 

5.70 

7.24 

5.96 

3.11 

3.35 

5.22 

291 

7.56 

47.24 

3.73 

4.66 

6.29 

5.20 

2.67 

3.15 

4.61 

292 

8.03 

41.48 

3.46 

4.35 

5.67 

5.60 

2.82 

2.94 

4.31 

293 

8.54 

44.54 

3.26 

4.93 

5.98 

5.65 

2.75 

2.95 

4.84 

294 

8.15 

42.02 

3.27 

3.93 

5.35 

5.32 

2.87 

2.71 

4.08 

295 

9.02 

50.62 

3.56 

4.64 

6.05 

5.84 

2.90 

2.97 

4.27 

296 

6.90 

43.50 

3.09 

4.22 

5.69 

4.89 

2.74 

2.48 

4.28 

297 

7.12 

39.29 

3.62 

4.49 

5.61 

5.18 

2.81 

2.98 

4.27 

298 

8.04 

42.24 

2.96 

4.30 

5.33 

5.45 

2.77 

2.52 

4.01 

299 

7.44 

43.89 

3.60 

4.49 

5.55 

5.68 

3.04 

2.87 

4.49 

300 

6.57 

38.10 

3.11 

3.70 

4.73 

4.22 

2.38 

2.51 

3.51 

301 

6.90 

38.99 

3.01 

3.86 

4.94 

4.57 

2.55 

2.84 

3.92 

302 

7.14 

36.36 

3.32 

3.85 

4.94 

4.36 

2.41 

2.75 

3.64 

303 

6.57 

37.42 

3.06 

3.84 

5.23 

4.50 

2.57 

2.60 

3.56 

304 

5.68 

31.24 

2.60 

3.26 

4.27 

3.78 

1.99 

1.97 

2.99 

305 

7.40 

34.73 

2.73 

3.52 

4.43 

3.67 

2.20 

2.17 

3.21 

306 

5.65 

32.04 

2.65 

3.31 

4.35 

3.84 

2.24 

2.21 

3.30 

307 

5.85 

36.06 

2.35 

2.52 

3.55 

2.73 

1.44 

1.72 

2.38 

308 

6.70 

33.52 

2.37 

2.99 

4.27 

3.66 

1.94 

2.04 

2.79 

309 

7.39 

37.41 

3.34 

4.11 

5.39 

4.48 

2.62 

2.55 

3.98 

310 

7.03 

35.55 

2.84 

3.54 

4.52 

3.90 

2.19 

2.29 

3.39 

311 

5.37 

35.21 

2.54 

3.47 

4.62 

3.87 

2.07 

2.12 

3.18 

312 

7.78 

38.16 

3.02 

3.75 

4.81 

4.02 

2.30 

2.41 

3.49 

313 

7.11 

38.56 

2.92 

3.73 

4.61 

2.25 

2.38 

2.41 

3.54 

314 

7.96 

43.28 

3.45 

4.15 

5.23 

4.90 

2.61 

2.76 

4.21 

315 

6.90 

30.73 

2.17 

2.94 

4.13 

3.60 

2.07 

1.91 

2.82 

316 

5.73 

30.75 

2.69 

3.34 

4.25 

3.56 

2.06 

2.13 

3.23 

317 

• 

37.86 

3.21 

4.10 

5.01 

4.95 

2.94 

2.70 

3.89 

318 

6.16 

32.91 

2.27 

2.95 

4.12 

3.18 

1.68 

1.68 

2.69 

319 

6.53 

29.76 

2.10 

2.61 

3.83 

3.29 

1.34 

1.71 

2.29 

320 

5.09 

28.95 

2.38 

2.97 

3.92 

3.37 

2.01 

2.07 

2.89 

321 

5.13 

27.52 

2.74 

3.22 

4.24 

3.49 

1.97 

2.09 

3.17 

322 

5.40 

34.31 

3.05 

3.63 

4.71 

4.15 

2.44 

2.41 

3.62 

323 

5.64 

26.81 

2.17 

2.68 

3.77 

3.20 

1.83 

1.81 

2.62 

324 

6.18 

29.69 

2.32 

2.81 

3.70 

3.14 

1.81 

1.84 

2.69 

325 

4.26 

24.68 

2.26 

2.72 

3.55 

3.44 

2.14 

1.97 

2.80 

326 

3.67 

22.94 

1.77 

2.46 

3.21 

2.97 

1.73 

1.55 

2.17 

327 

4.44 

22.92 

1.95 

2.46 

2.94 

2.74 

1.43 

1.59 

2.17 

328 

3.92 

21.74 

1.79 

2.27 

2.94 

2.67 

1.36 

1.42 

2.16 

329 

• 

20.77 

2.02 

2.41 

2.85 

2.83 

1.69 

1.63 

2.29 

330 

3.37 

20.44 

1.65 

1.95 

2.74 

2.58 

1.52 

1.35 

1.82 

331 

3.96 

20.54 

1.98 

2.31 

2.84 

2.85 

1.68 

1.65 

2.14 

332 

• 

19.18 

1.82 

1.91 

2.73 

2.57 

1.59 

1.41 

1.83 

333 

• 

19.09 

1.67 

1.87 

2.68 

2.57 

1.42 

1.34 

1.80 

334 

16.12 

101.02 

6.10 

8.19 

• 

10.94 

4.50 

4.55 

7.92 

335 

9.98 

67.17 

4.77 

5.94 

7.57 

8.15 

3.38 

3.37 

6.16 

336 

10.45 

57.62 

4.24 

6.09 

8.60 

8.28 

3.84 

3.39 

5.42 


380 


# 

PDV 

TWMX 

LTH1 

LTH2 

LTH3 

LTH4 

LTH5 

TTH1 

TTH2 

337 

9.49 

51.90 

3.92 

4.99 

6.77 

6.98 

3.41 

3.35 

4.88 

338 

8,87 

49.92 

3,44 

4.87 

6.52 

5.61 

2.79 

2.68 

4.36 

339 

9.15 

47.59 

3.91 

5.55 

7.03 

7.18 

3.77 

3.23 

5.48 

340 

7.52 

45.67 

3.08 

4.03 

5.94 

5.26 

2.33 

2.47 

3.81 

341 

7.93 

44.80 

3.81 

4.86 

6.15 

5.33 

3.47 

3.09 

4,86 

342 

7,88 

46.74 

2.11 

3.11 

4.65 

3.09 

1.52 

1.74 

2.38 

343 

8.26 

43.09 

3.09 

4.02 

5.55 

5,01 

2.16 

2.58 

3.89 

344 

6.62 

38.72 

3.93 

4.49 

5.49 

5.31 

3.23 

3.15 

4.32 

345 

6.96 

37.91 

2.93 

3.82 

4.81 

4.34 

2.33 

2,45 

3.66 

346 

6.04 

36.31 

2.84 

3.74 

4.64 

3.83 

2.03 

2.18 

3.15 

347 

7.11 

31.69 

2.64 

3.18 

4.48 

4.24 

2.18 

2.18 

3.10 

348 

5.43 

25.81 

2.38 

2.61 

3.61 

3.10 

2.09 

1.97 

2,42 

349 

• 

24.81 

1.84 

2.43 

3.30 

3.22 

1.57 

1,78 

2.20 

350 

3.95 

24,76 

2.47 

2.65 

3.69 

3.11 

2.08 

2.05 

2.63 

351 

4.57 

23.56 

2.09 

2.39 

3.21 

2.91 

1.60 

1.56 

2.10 

352 

3.61 

19.72 

2.03 

2.27 

3.04 

3.18 

1.98 

1.89 

2.28 

353 

• 

15.05 

1.63 

1.88 

2.52 

2.16 

1,36 

1.34 

1.92 

354 

12.06 

66.76 

• 

9.11 

9.65 

• 

1.67 

1.76 

6.46 

355 

15,04 

83.71 

3.13 

• 


• 

2.09 

2.60 

• 

356 


88.86 

3.92 

8.92 


• 

2.19 

2.67 

• 

357 

16.72 

84.39 


• 


• 

2.24 

3.00 

• 

358 

• 

• 

• 

• 

• 


• 

3.36 

• 

359 

15.35 

89.02 

3.00 

• 

• 

• 

2.65 

3.06 

8.96 

360 

• 

94.47 

• 

• 

• 

• 

• 

3.31 

• 

361 

• 

• 

4.49 

• 


10.80 

3.69 

• 

• 

362 

12.45 

66.73 

2.70 

6.61 

8.29 

6.70 

2,34 

2.70 

6.71 

363 

• 

66.09 

2.37 

• 

• 

6.93 

1.85 

2.22 

• 

364 

• 

70.98 

« 

• 

• 

• 

• 

• 

• 

365 

7.37 

62.19 

• 

« 

• 

• 

• 

• 

• 

366 

• 

28.74 

• 

• 

• 

• 

• 

• 

• 

367 

• 

25.84 

• 

« 

• 

• 

• 

• 

• 

368 

• 

25.18 

• 

• 

• 

• 

• 

• 

• 

369 

• 

24,28 

• 

• 

• 

• 

• 

• 

• 

370 

10.52 

77.93 

• 

• 

• 

• 

• 

• 

• 

371 

• 

90.39 

3,36 

9.37 

• 

9.81 

1.81 

• 

11.12 

372 

22.03 

139.81 

3.32 

8.07 

• 

8.80 

1.82 

1.81 

7.83 


# 

1 

2 

3 

4 

5 

6 

7 

8 

9 

10 

11 

12 

13 

14 

15 

16 

17 

18 

19 

20 

21 

22 

23 

24 

25 

26 

27 

28 

29 

30 

31 

32 

33 

34 

35 

36 

37 

38 

39 

40 

41 

42 

43 

44 

45 

46 

47 

48 


381 


TTH3 

TTH4 

TTH5 

PALI 

PALM 

PALIII 

PALIV 

PALV 

7.74 

5.56 

1.87 

25.60 

18.52 

17.38 

18.05 

25.10 

5.13 

3.11 

1.30 

12.17 

9.09 

10.16 

9.67 

12.36 

3.72 

2.03 

1.06 

9.44 

7.03 

7.45 

7.16 

9.28 

3.42 

2.26 

1.13 

9.32 

6.41 

7.59 

7.13 

8.52 

3.25 

1.86 

0.88 

6.55 

4.82 

5.46 

5.23 

6.46 

2.57 

1.82 

0.97 

4.92 

4.26 

4.97 

4.68 

5.43 

6.75 

5.01 

1.57 

23.77 

17.62 

17.22 

16.64 

23.36 

4.47 

2.94 

1.29 

12.80 

9.35 

9.49 

9.82 

12.78 

3.91 

2.85 

1.38 

12.47 

10.08 

9.80 

9.98 

11.83 

5.20 

2.99 

1.66 

11.72 

8.77 

9.63 

10.04 

12.78 

4.58 

2.54 

1.33 

10.73 

7.87 

8.53 

8.36 

10.99 

3.88 

2.48 

0.95 

10.66 

7.32 

8.48 

7.84 

10.79 

3.54 

2.31 

1.07 

7.11 

6.04 

6.01 

• 

7.53 

3.27 

1.63 

1.00 

7.18 

5.13 

6.32 

5.91 

7.06 

• 

1.66 

0.83 

• 

• 

• 

4.78 

5.57 

3.49 

2.31 

1.06 

10.47 

8.52 

8.86 

8.58 

10.52 

7.73 

5.70 

1.87 

25.46 

17.38 

16.45 

17.44 

24.65 

6.60 

4.80 

1.62 

18.97 

14.83 

14.42 

14.73 

19.35 

6.01 

3.86 

• 

16.81 

12.64 

13.10 

12.62 

16.61 

8.05 

5.54 

1.48 

24.44 

17.47 

• 

18.05 

23.67 

3.78 

2.20 

1.14 

8.54 

• 

• 

• 

• 

3.37 

1.84 

1.14 

• 

• 

• 

• 

• 

3.89 

2.45 

1.38 

9.56 

7.12 

8.01 

7.27 

• 

3.69 

2.15 

0.95 

9.53 

6.37 

• 

6.76 

• 

3.64 

2.47 

• 

10.68 

8.59 

• 

8.92 

12.09 

3.81 

1.94 

0.93 

8.42 

6.00 

7.15 

6.71 

8.50 

3.26 

1.96 

0.97 

8.57 

6.48 

7.40 

6.84 

8.52 

5.77 

3.39 

1.38 

15.40 

11.88 

12.39 

11.59 

15.08 

4.72 

2.88 

1.34 

10.24 

• 

• 

• 

10.60 

3.53 

2.08 

1.09 

• 

• 

• 

• 

• 

2.84 

1.70 

0.78 

• 

• 

• 

• 

• 

2.81 

1.53 

0.83 

5.95 

4.85 

5.14 

5.23 

6.41 

2.20 

1.23 

0.76 

• 

• 

• 

• 

• 

2.15 

1.38 

0.93 

• 

• 

• 

• 

• 

6.89 

4.59 

1.14 

20.96 

13.80 

13.88 

14.38 

21.25 

5.79 

4.00 

1.38 

17.15 

13.01 

13.83 

• 

17.47 

6.52 

4.61 

1.46 

20.26 

14.82 

15.25 

15.31 

19.87 

3.41 

1.79 

0.73 

8.08 

• 

• 

6.17 

8.47 

3.73 

2.13 

1.07 

• 

• 

• 

• 

• 

3.56 

2.31 

1.00 

8.92 

6.76 

7.31 

7.06 

9.00 

3.00 

1.81 

0.93 

• 

• 

7.49 

7.27 

8.82 

3.54 

2.17 

1.07 

8.25 

6.05 

6.62 

6.52 

8.34 

3.47 

1.82 

0.86 

• 

• 

5.76 

5.39 

6.94 

3.28 

1.89 

1.06 

6.75 

5.15 

• 

• 

6.90 

2.94 

1.99 

0.86 

5.95 

• 

4.59 

4.52 

6.17 

• 

4.55 

1.68 

• 

• 

• 

• 

• 

3.58 

1.89 

1.00 

• 

5.37 

• 

• 

• 

3.08 

1.81 

0.90 

7.39 

5.28 

• 

5.98 

7.35 


382 


# 

TTH3 

TTH4 

TTH5 

PALI 

PALII 

PALIN 

PALIV 

PALV 

49 

2.40 

1.42 

0.71 

5.51 

4.97 

• 

5.09 

5.63 

50 

2.76 

1.73 

0.97 

7.25 

5.82 

5.99 

6.00 

• 

51 

2.42 

1.71 

0.73 

• 

4.91 

5.25 

4.88 

6.45 

52 

2.50 

1.79 

0.91 

• 

• 

• 

5.13 

6.17 

53 

2.27 

1.62 

0.95 

• 

4.55 

• 

4.58 

• 

54 

2.22 

1.53 

0.88 

• 

• 

• 

• 

5.77 

55 

2.04 

1.24 

0.71 

4.72 

3.93 

4.45 

4.12 

5.20 

56 

2.10 

1.28 

0.68 

• 

• 

• 

3.63 

4.19 

57 

1.65 

0.96 

0.67 

3.23 

• 

3.26 

3.16 

• 

58 

5.03 

2.84 

1.45 

12.68 

8.94 

9.86 

9.48 

12.57 

59 

4.29 

2.20 

1.25 

• 

• 

8.52 

8.93 

10.81 

60 

6.14 

3.43 

1.57 

15.59 

11.85 

12.63 

12.37 

16.12 

61 

4.54 

2.63 

1.26 

10.94 

8.39 

8.82 

8.54 

10.97 

62 

• 

2.34 

1.21 

7.89 

6.38 

6.81 

6.77 

7.76 

63 

4.68 

2.92 

1.36 

11.24 

9.24 

9.91 

9.87 

11.47 

64 

4.04 

2.47 

1.19 

9.34 

• 

6.73 

6.71 

9.31 

65 

3.91 

2.18 

1.26 

9.16 

6.78 

7.30 

7.14 

9.22 

66 

7.13 

5.13 

1.93 

24.05 

18.51 

19.22 

19.06 

25.12 

67 

3.59 

2.47 

1.08 

11.17 

8.25 

8.93 

8.36 

11.22 

68 

• 

2.79 

1.53 

12.23 

8.76 

9.34 

8.98 

12.36 

69 

5.96 

3.74 

1.55 

14.92 

10.44 

11.60 

11.15 

15.17 

70 

3.80 

2.61 

1.28 

• 

• 

• 

• 

• 

71 

4.79 

3.06 

1.34 

• 

• 

• 

• 

• 

72 

3.23 

2.19 

1.19 

8.48 

6.50 

7.20 

6.88 

8.70 

73 

6.12 

4.23 

1.61 

19.09 

13.82 

13.87 

14.13 

18.99 

74 

4.14 

2.50 

1.42 

• 

• 

• 

• 

• 

75 

3.13 

2.16 

1.16 

9.94 

7.79 

8.84 

8.71 

10.09 

76 

• 

• 

• 

12.93 

9.79 

9.95 

• 

13.79 

77 

• 

5.58 

1.83 

• 

• 

• 

19.04 

27.49 

78 

2.91 

1.68 

0.93 

6.59 

5.49 

• 

5.67 

6.95 

79 

3.53 

2.09 

1.15 

8.54 

6.24 

7.01 

• 

8.63 

80 

3.28 

1.99 

1.04 

7.20 

5.30 

6.18 

5.74 

• 

81 

2.61 

1.45 

1.14 

5.74 

• 

• 

• 

• 

82 

3.15 

2.06 

1.21 

• 

• 

• 

6.77 

8.65 

83 

• 

1.49 

0.93 

6.07 

4.58 

5.31 

4.80 

• 

84 

3.51 

2.35 

1.18 

• 

• 

• 

• 

• 

85 

• 

• 

• 

10.23 

• 

• 

8.25 

• 

86 

• 

• 

• 

• 

7.53 

7.32 

• 

9.98 

87 

3.79 

2.16 

1.21 

9.57 

7.32 

7.53 

7.47 

9.55 

88 

4.77 

2.96 

1.48 

12.67 

9.67 

10.16 

9.91 

12.91 

89 

7.85 

5.14 

1.79 

25.91 

18.86 

• 

19.52 

• 

90 

• 

• 

• 

6.45 

5.17 

6.35 

6.01 

6.89 

91 

3.29 

1.93 

1.08 

• 

• 

6.59 

• 

• 

92 

2.91 

1.64 

0.95 

• 

• 

. 

• 

• 

93 

2.36 

1.39 

0.88 

5.87 

4.83 

5.29 

• 

• 

94 

2.97 

1.81 

1.01 

• 

5.11 

6.37 

• 

• 

95 

3.11 

1.97 

1.11 

7.30 

5.31 

6.51 

6.15 

7.64 

96 

2.99 

1.83 

1.14 

• 

5.59 

6.05 

5.70 

• 


383 


# 

TTH3 

TTH4 

TTH5 

PALI 

PALII 

PALI II 

PALIV 

PALV 

97 

2.93 

1.56 

1.07 

• 

• 

• 

• 

• 

98 

8.29 

5.46 

1.83 

25.95 

19.61 

19.12 

19.93 

• 

99 

6.28 

4.09 

1.58 

18.42 

13.66 

12.96 

13.48 

17.92 

100 

7.20 

4.58 

1.50 

22.36 

16.72 

16.85 

17.28 

22.90 

101 

• 

3.51 

1.53 

• 

• 

• 

• 

• 

102 

5.91 

3.42 

1.35 

18.67 

13.99 

14.89 

14.47 

19.45 

103 

3.41 

1.88 

1.05 

8.70 

6.27 

• 

6.18 

• 

104 

2.56 

1.81 

0.94 

7.24 

• 

• 

• 

• 

105 

3.43 

2.11 

0.99 

7.41 

5.67 

• 

• 

7.30 

106 

3.07 

1.86 

1.10 

• 

4.93 

• 

4.89 

• 

107 

3.28 

1.83 

1.15 

• 

• 

• 

• 

• 

108 

3.05 

2.21 

1.15 

• 

5.22 

• 

5.57 

• 

109 

2.76 

1.77 

0.96 

6.61 

5.44 

6.08 

5.76 

6.97 

110 

3.62 

2.22 

1.35 

• 

• 

• 

• 

8.58 

111 

3.05 

1.87 

1.16 

7.04 

• 

• 

5.41 

6.92 

112 

2.62 

1.51 

0.90 

5.76 

4.17 

4.63 

4.48 

6.01 

113 

2.53 

1.69 

0.89 

6.12 

5.07 

• 

5.28 

• 

114 

3.37 

1.82 

0.98 

• 

• 

• 

• 

• 

115 

2.48 

1.79 

0.94 

5.56 

4.66 

• 

• 

• 

116 

3.24 

1.79 

1.22 

7.67 

• 

• 

• 

• 

117 

2.31 

1.41 

0.89 

5.04 

4.41 

• 

4.20 

5.07 

118 

2.13 

1.29 

0.87 

• 

3.15 

3.66 

3.60 

• 

119 

2.16 

1.46 

0.99 

• 

• 

• 

• 

• 

120 

7.01 

4.39 

1.74 

21.72 

16.32 

16.12 

17.15 

21.76 

121 

5.19 

2.88 

1.44 

13.52 

10.13 

10.64 

11.03 

14.30 

122 

3.07 

2.04 

1.02 

• 

5.52 

• 

• 

7.71 

123 

2.54 

1.67 

0.95 

• 

• 

• 

5.19 

• 

124 

2.75 

1.96 

1.15 

6.82 

• 

5.50 

5.64 

7.14 

125 

2.70 

1.58 

1.01 

6.39 

4.94 

5.53 

5.05 

6.21 

126 

2.29 

1.49 

0.87 

• 

• 

• 

4.05 

• 

127 

• 

5.41 

1.79 

• 

• 

• 

• 

• 

128 

4.03 

2.92 

1.26 

10.59 

8.00 

8.41 

9.01 

10.45 

129 

7.80 

4.83 

1.78 

23.75 

17.59 

• 

• 

• 

130 

• 

6.81 

1.79 

• 

21.84 

21.84 

23.21 

23.95 

131 

7.78 

6.09 

1.28 

17.47 

13.44 

15.11 

14.94 

18.78 

132 

8.89 

7.20 

2.85 

20.62 

• 

• 

17.13 

• 

133 

8.24 

6.19 

2.46 

12.45 

11.11 

12.34 

11.49 

13.72 

134 

5.25 

3.46 

1.42 

15.64 

10.57 

10.27 

10.46 

13.74 

135 

8.71 

6.22 

2.59 

15.62 

13.60 

13.90 

13.38 

16.43 

136 

• 

5.48 

2.03 

• 

14.09 

15.03 

14.02 

« 

137 

2.55 

1.80 

0.96 

5.66 

4.10 

4.95 

4.61 

5.71 

138 

8.00 

5.94 

2.46 

19.88 

13.98 

15.67 

• 

• 

139 

7.50 

5.28 

2.08 

17.48 

13.56 

15.15 

14.42 

16.30 

140 

6.99 

4.92 

2.08 

14.42 

11.58 

12.72 

12.48 

14.79 

141 

2.73 

2.15 

1.16 

• 

• 

• 

• 

• 

142 

8.43 

6.31 

2.89 

18.28 

14.32 

15.08 

14.77 

17.88 

143 

• 

7.53 

2.55 

• 

• 

• 

• 

• 

144 

2.59 

1.82 

1.05 

5.16 

4.67 

4.41 

4.52 

5.35 


384 


# 

TTH3 

TTH4 

TTH5 

PALI 

PALM 

PALIII 

PALIV 

PALV 

145 

3.40 

2.33 

1.28 

8.02 

6.66 

7.73 

7.77 

8.68 

146 

• 

2.57 

1.28 

• 

• 

• 

• 

• 

147 

9.86 

8.25 

3.13 

22.21 

17.65 

17.91 

17.67 

21.48 

148 

8.33 

5.85 

2.42 

18.79 

13.75 

15.73 

12.95 

18.75 

149 

• 

5.58 

1.52 

34.67 

22.65 

27.27 

24.85 

34.48 

150 

• 

5.85 

1.42 

27.68 

18.78 

23.66 

20.19 

28.11 

151 

• 

4.49 

1.27 

26.93 

18.92 

22.16 

20.26 

28.95 

152 

• 

6.61 

1.53 

36.04 

25.51 

31.03 

26.08 

36.06 

153 

• 

4.35 

1.36 

21.18 

15.98 

19.97 

16.90 

21.68 

154 

• 

5.66 

1.38 

27.68 

20.15 

22.09 

20.05 

27.60 

155 

• 

7.75 

1.69 

• 

• 

• 

24.96 

33.30 

156 

7.79 

• 

1.07 

• 

16.93 

19.04 

17.50 

22.29 

157 

9.13 

4.38 

0.91 

27.78 

18.93 

22.83 

19.24 

27.48 

158 


5.33 

1.28 

31.15 

22.61 

27.38 

23.73 

30.70 

159 


4.49 

1.14 

22.45 

16.54 

19.38 

17.60 

22.97 

160 

• 

6.51 

1.54 

35.17 

25.62 

29.78 

26.78 

34.55 

161 

• 

5.33 

1.47 

29.14 

21.94 

25.88 

23.11 

29.95 

162 

• 

4.78 

1.39 

25.86 

20.02 

22.09 

19.82 

25.28 

163 


4.61 

1.16 

23.30 

16.80 

19.82 

17.43 

22.99 

164 

• 

4.36 

1.20 

23.68 

17.81 

20.56 

18.05 

23.72 

165 


7.31 

1.71 

34.92 

26.81 

27.62 

• 

35.10 

166 

• 

• 

• 

37.51 

27.69 

33.07 

28.47 

37.15 

167 

• 

4.40 

1.16 

26.12 

19.76 

24.53 

19.64 

26.33 

168 

9.52 

5.43 

0.99 

25.36 

17.17 

21.03 

19.57 

25.48 

169 

• 

4.08 

1.11 

23.26 

17.52 

22.08 

20.75 

23.69 

170 

• 

7.31 

• 

23.17 

16.86 

22.15 

18.12 

23.53 

171 

8.63 

4.10 

1.22 

22.57 

16.82 

19.41 

• 

22.63 

172 

• 

6.41 

1.72 

31.12 

22.54 

24.80 

23.07 

32.70 

173 

• 

6.35 

1.68 

29.48 

23.08 

24.39 

23.67 

30.56 

174 

• 

5.57 

1.12 

26.47 

18.41 

20.99 

18.19 

26.05 

175 

8.52 

3.88 

1.01 

21.66 

15.50 

18.44 

• 

21.06 

176 

• 

6.37 

1.39 

33.57 

24.92 

27.01 

24.69 

34.27 

177 

8.76 

4.41 

1.15 

21.96 

16.74 

19.60 

17.07 

22.87 

178 

10.27 

• 

• 

28.97 

19.96 

• 

20.06 

28.90 

179 

• 

5.37 

1.15 

29.10 

22.63 

23.44 

23.79 

29.67 

180 

• 

4.81 

1.15 

29.08 

20.39 

22.84 

21.19 

28.33 

181 

9.42 

• 

1.13 

23.81 

16.51 

20.36 

17.30 

24.75 

182 

• 

4.23 

1.28 

25.87 

18.51 

21.89 

19.62 

26.59 

183 

• 

4.50 

1.25 

29.68 

21.49 

24.89 

22.89 

30.24 

184 

• 

4.64 

1.34 

26.35 

19.16 

23.22 

20.24 

26.31 

185 

• 

6.89 

1.63 

32.58 

22.47 

26.23 

24.50 

32.67 

186 

• 

3.71 

1.32 

23.19 

16.91 

22.03 

18.88 

23.95 

187 

• 

5.97 

1.51 

28.98 

22.01 

24.81 

21.92 

30.02 

188 

• 

7.13 

1.76 

32.53 

22.88 

28.74 

24.23 

33.55 

189 

• 

5.28 

1.35 

30.27 

22.64 

26.35 

22.86 

30.49 

190 

• 

5.23 

1.51 

27.20 

18.82 

21.03 

18.48 

27.01 

191 

• 

4.59 

1.19 

23.53 

18.75 

22.25 

19.74 

25.79 

192 

• 

4.19 

1.09 

26.94 

18.09 

23.06 

19.27 

26.92 


385 


# 

TTH3 

TTH4 

TTH5 

PALI 

193 

• 

5.29 

1.18 

28.02 

194 

7.21 

4.48 

0.83 

16.87 

195 

• 

5.09 

1.47 

25.32 

196 

• 

4.34 

0.81 

23.34 

197 

• 

4.82 

1.38 

29.84 

198 

• 

4.03 

0.83 

25.78 

199 

• 

5.34 

1.43 

25.14 

200 

• 

4.58 

1.21 

26.18 

201 

• 

4.53 

1.42 

26.40 

202 

• 

5.06 

1.28 

29.23 

203 

7.52 

3.85 

1.27 

17.86 

204 

• 

5.89 

1.43 

34.23 

205 

6.92 

3.22 

1.07 

17.18 

206 


8.77 

1.73 

37.74 

207 

• 

4.52 

1.18 

24.04 

208 

9.03 

4.17 

1.46 

23.43 

209 

5.85 

2.87 

• 

16.24 

210 

7.69 

3.54 

1.15 

18.81 

211 

• 

4.11 

1.17 

25.37 

212 


4.76 

1.20 

26.78 

213 

• 

5.38 

1.47 

28.90 

214 

• 

6.28 

1.54 

29.61 

215 

• 

5.64 

1.27 

26.33 

216 


4.99 

1.30 

34.84 

217 


4.27 

1.34 

26.21 

218 

• 

5.62 

1.26 

28.19 

219 

• 

5.13 

1.52 

29.39 

220 

• 

4.49 

1.35 

24.92 

221 


4.86 

1.26 

25.18 

222 


6.58 

1.48 

31.46 

223 


7.49 

1.57 

25.47 

224 


8.21 

1.93 

36.19 

225 


8.31 

1.52 

27.48 

226 

• 

6.64 

1.67 

17.89 

227 

• 

10.43 


• 

228 

• 

8.33 

1.86 

34.44 

229 

• 

9.14 

1.95 

33.12 

230 

• 

• 

1.72 

24.18 

231 

• 

7.13 

1.94 

36.65 

232 

• 

6.54 

1.38 

24.86 

233 

8.64 

5.82 

3.68 

32.27 

234 

• 

6.46 

3.22 

34.22 

235 

9.43 

6.47 

2.81 

34.53 

236 

10.70 

7.91 

3.74 

• 

237 

7.93 

6.14 

3.69 

29.54 

238 

8.12 

5.80 

2.95 

29.43 

239 

7.62 

6.08 

3.49 

19.93 

240 

7.62 

6.24 

2.95 

26.08 


PALM 

PALIN 

PALIV 

PALV 

21.19 

23.64 

21.76 

27.85 

13.13 

16.54 

13.69 

17.41 

18.72 

22.55 

19.71 

26.77 

16.73 

20.32 

18.45 

24.65 

19.87 

24.90 

20.12 

28.90 

18.18 

20.20 

18.47 

25.05 

18.71 

20.81 

19.10 

25.84 

19.51 

22.28 

19.62 

26.54 

19.64 

23.46 

19.92 

26.32 

21.72 

25.29 

23.57 

29.95 

14.06 

17.37 

14.96 

18.58 

25.90 

28.70 

25.27 

34.07 

13.13 

15.64 

14.03 

17.32 

27.94 

29.52 

28.39 

37.54 

16.56 

21.43 

17.61 

24.54 

17.92 

21.98 

19.28 

24.56 

13.21 

15.94 

13.03 

16.40 

14.04 

16.76 

14.69 

19.95 

17.26 

20.77 

18.07 

25.64 

• 

21.78 

19,76 

• 

20.14 

23.90 

21.37 

29.05 

19.77 

24.10 

20.03 

29.86 

19.83 

• 

22.47 

27.15 

22.10 

26.03 

23.48 

34.49 

19.07 

23.42 

20.55 

26.80 

19.98 

24.52 

21.70 

28.69 

20.98 

25.83 

21.77 

29.71 

18.62 

22.10 

19.32 

24.94 

18.89 

21.86 

19.30 

25.58 

24.67 

26.57 

24.25 

32.30 

18.77 

23.02 

• 

27.16 

30.55 

32.76 

32.58 

37.56 

21.57 

23.16 

22.44 

28.06 

14.17 

16.71 

15.18 

18.47 

26.26 

29.93 

27.87 

35.87 

25.93 

28.43 

26.31 

34.69 

18.75 

22.58 

19.25 

24.70 

26.96 

29.87 

27.01 

36.52 

18.36 

22.27 

18.99 

26.03 

22.98 

26.51 

24.42 

32.49 

26.63 

• 

26.92 

• 

26.09 

28.15 

26.22 

35.25 

25.42 

27.95 

27.01 

34.15 

• 

• 

21.66 

29.57 

20.75 

20.61 

20.67 

• 

• 

15.81 

16.23 

• 

19.98 

21.10 

20.01 

27.13 


386 


# 

TTH 3 

TTH 4 

241 

6.95 

5.15 

242 

5.91 

4.38 

243 

5.91 

4.40 

244 

6.04 

4.52 

245 

5.88 

4.61 

246 

4.94 

3.61 

247 

5.49 

4.50 

248 

5.35 

4.34 

249 

4.76 

3.42 

250 

4.44 

3.41 

251 

5.30 

3.93 

252 

4.72 

3.35 

253 

4.19 

3.11 

254 

3.31 

2.31 

255 

4.72 

3.55 

256 

4.27 

3.04 

257 

4.25 

3.27 

258 

3.98 

3.23 

259 

3.83 

2.57 

260 

3.30 

2.28 

261 

3.59 

2.57 

262 

2.84 

2.10 

263 

9.54 

6.33 

264 

9.14 

6.77 

265 

6.53 

5.09 

266 

• 

9.77 

267 

• 

8.68 

268 

• 

9.07 

269 

• 

7.39 

270 

• 

7.42 

271 

• 

7.81 

272 

9.41 

6.73 

273 

7.45 

6.05 

274 

7.39 

6.76 

275 

9.04 

6.73 

276 

5.65 

4.23 

277 

8.53 

6.71 

278 

8.16 

6.06 

279 

5.84 

4.78 

280 

7.96 

5.31 

281 

6.18 

4.57 

282 

7.83 

5.17 

283 

7.56 

4.94 

284 

7.52 

5.45 

285 

6.11 

4.49 

286 

7.01 

4.72 

287 

6.92 

4.85 

288 

6.21 

4.69 


TTH 5 

PALI 

PALII 

3.06 

22.21 

14.63 

2.23 

19.71 

13.61 

1.98 

16.33 

11.76 

2.31 

14.09 

11.37 

2.56 

13.28 

10.43 

1.65 

13.95 

10.42 

2.59 

13.34 

9.77 

2.61 

13.56 

9.79 

2.00 

13.27 

• 

2.18 

12.61 

9.32 

2.42 

12.43 

9.37 

2.15 

• 

8.82 

1.87 

11.88 

8.53 

1.47 

• 

6.28 

2.13 

• 

• 

1.94 

• 

• 

2.00 

11.04 

8.85 

2.14 

10.92 

7.78 

1.71 

10.51 

7.95 

1.54 

• 

• 

1.63 

9.62 

7.07 

1.25 

6.62 

5.25 

4.04 

24.99 

19.16 

3.51 

24.76 

19.57 

3.28 

18.81 

13.73 

4.17 

37.35 

30.80 

4.44 

37.05 

27.78 

4.35 

39.79 

30.53 

4.55 

37.37 

28.32 

4.03 

• 

26.06 

4.44 

32.08 

26.85 

3.83 

29.13 

21.15 

3.96 

24.89 

18.42 

3.95 

20.55 

15.32 

4.17 

22.40 

16.31 

2.46 

15.36 

11.82 

4.31 

23.78 

18.34 

3.88 

21.31 

16.71 

3.14 

17.76 

12.28 

3.39 

26.26 

20.19 

3.22 

19.74 

15.01 

3.25 

26.17 

18.26 

2.98 

22.22 

16.46 

3.57 

• 

17.75 

3.15 

15.94 

12.26 

3.27 

22.54 

16.11 

3.07 

19.35 

13.83 

2.95 

15.92 

12.45 


PALIN 

PALIV 

PALV 

16.52 

15,82 

21.90 

14.50 

13.99 

19.78 

13.91 

12.17 

16.37 

12.80 

12,02 

14.83 

11.03 

10.74 

13.79 

11.50 

10.98 

14.54 

10.51 

10,54 

13.71 

11.15 

10.21 

14,23 

10.31 

10.14 

13.86 

• 

9.89 

13.86 

11.18 

9.69 

13.28 

9.70 

9.44 

12.67 

6.38 

• 

• 

• 

• 

• 

• 

9.24 

• 

9.11 

• 

11.39 

8.34 

8.39 

11.36 

7.72 

7.31 

9.14 

5.27 

5.39 

6.51 

19.77 

20.11 

25.96 

19.02 

19.73 

25.83 

14.70 

15.09 

20.09 

32.58 

30.53 

38.18 

28.86 

26.63 

37.82 

33.19 

30.26 

41.86 

33.22 

30.62 

38.53 

28.41 

26.11 

• 

22.02 

20.37 

29.17 

20.60 

19.03 

25.51 

15.84 

16.29 

22.36 

18.58 

18.52 

23.51 

11.89 

12.15 

15.25 

19.98 

19.48 

25.11 

17.59 

16,95 

22.47 

12.83 

12.58 

18.23 

21.02 

20.41 

26.24 

15.77 

15.55 

20.91 

19.55 

• 

27.02 

16.96 

17.29 

24,12 

18.57 

17.36 

22.12 

13.02 

12.80 

17.44 

16.63 

17.38 

23,25 

16.03 

14.85 

20.22 

14.46 

12.93 

16.33 


387 


# 

TTH3 

TTH4 

TTH5 

PALI 

PALM 

PALIN 

PALIV 

PALV 

289 

5.59 

4.13 

2.66 

15.17 

12.52 

• 

• 

15.92 

290 

7.25 

5.26 

3.30 

21.29 

15.04 

15.09 

15.48 

21.62 

291 

6.23 

4.61 

3.25 

17.32 

12.36 

12.79 

12.72 

17.71 

292 

5.69 

4.04 

2.94 

12.61 

10.83 

11.45 

11.30 

13.70 

293 

5.98 

4.71 

2.94 

17.96 

12.29 

12.13 

12.37 

17.95 

294 

5.37 

3.92 

2.86 

14.36 

10.63 

11.88 

11.54 

15.24 

295 

6.08 

4.55 

3.01 

16.93 

13.01 

14.43 

13.94 

17.93 

296 

5.67 

4.01 

2.72 

15.30 

12.31 

13.55 

12.59 

• 

297 

5.62 

4.44 

3.09 

13.75 

10.02 

11.23 

11.33 

14.88 

298 

5.27 

4.20 

2.75 

15.02 

11.70 

12.24 

11.50 

15.56 

299 

5.52 

4.42 

3.03 

15.21 

11.25 

12.25 

11.94 

16.62 

300 

4.67 

3.43 

2.61 

12.04 

• 

• 

10.46 

12.76 

301 

4.90 

3.71 

2.69 

13.26 

9.84 

• 

10.72 

14.20 

302 

4.91 

3.56 

2.64 

11.75 

8.80 

9.06 

9.02 

12.62 

303 

5.23 

3.58 

2.57 

13.75 

9.96 

10.79 

10.62 

13.61 

304 

4.30 

3.24 

2.18 

11.54 

8.43 

9.18 

8.89 

10.75 

305 

4.45 

3.06 

2.08 

12.82 

9.13 

9.72 

9.51 

12.62 

306 

4.35 

3.23 

2.23 

11.13 

8.43 

8.85 

8.66 

12.02 

307 

3.59 

2.53 

1.82 

9.77 

8.58 

8.92 

8.73 

10.59 

308 

4.24 

2.75 

1.86 

11.63 

9.17 

9.30 

8.94 

11.88 

309 

5.40 

3.84 

2.47 

13.35 

10.08 

10.84 

10.29 

13.86 

310 

4.47 

3.27 

2.33 

12.84 

9.47 

10.15 

9.14 

13.55 

311 

4.52 

3.12 

2.21 

10.61 

8.17 

9.05 

8.59 

11.75 

312 

4.82 

3.52 

2.27 

13.35 

10.13 

11.10 

10.99 

14.19 

313 

4.62 

3.43 

2.43 

12.67 

9.97 

11.40 

10.92 

14.27 

314 

5.19 

4.02 

2.92 

15.83 

12.32 

12.55 

12.07 

16.25 

315 

4.18 

2.86 

1.84 

11.34 

8.42 

9.05 

8.68 

11.89 

316 

4.22 

3.01 

2.30 

• 

• 

• 

8.62 

11.74 

317 

4.99 

3.89 

2.84 

12.84 

10.13 

10.11 

10.06 

13.30 

318 

4.08 

2.49 

1.82 

10.62 

• 

8.65 

9.68 

11.66 

319 

3.84 

2.20 

1.69 

9.79 

6.78 

7.72 

7.37 

9.67 

320 

3.94 

2.89 

1.94 

9.83 

7.40 

8.30 

7.98 

10.49 

321 

4.25 

3.14 

2.45 

• 

• 

7.37 

7.39 

9.51 

322 

4.74 

3.40 

2.43 

• 

9.04 

9.70 

• 

12.96 

323 

3.79 

2.66 

1.86 

9.19 

6.97 

7.53 

7.27 

9.74 

324 

3.72 

2.59 

1.90 

9.82 

6.98 

8.01 

7.50 

10.16 

325 

3.51 

2.70 

2.05 

8.42 

6.50 

• 

• 

9.11 

326 

3.19 

2.05 

1.53 

7.68 

6.23 

6.67 

6.58 

8.16 

327 

2.95 

2.14 

1.51 

7.01 

5.94 

• 

6.17 

7.76 

328 

2.97 

2.06 

1.42 

6.81 

5.78 

6.64 

• 

7.26 

329 

2.84 

2.22 

1.67 

• 

5.84 

• 

5.69 

7.58 

330 

2.73 

1.94 

1.45 

6.48 

• 

• 

5.37 

6.12 

331 

2.84 

2.26 

1.75 

5.91 

5.03 

• 

4.90 

6.02 

332 

2.69 

1.82 

1.40 

• 

• 

• 

4.87 

6.99 

333 

2.71 

1.88 

1.46 

5.76 

4.36 

• 

4.71 

6.33 

334 

• 

8.23 

4.80 

38.75 

29.54 

30.66 

30.48 

38.96 

335 

7.63 

6.18 

3.88 

24.27 

18.87 

• 

19.23 

25.12 

336 

8.62 

5.74 

3.42 

21.47 

15.47 

16.21 

15.83 

21.02 


388 


# 

TTH3 

TTH4 

TTH5 

PALI 

PALM 

PALIII 

PALIV 

PALV 

337 

6.76 

4.99 

3.46 

20.64 

15.17 

15.54 

15.49 

21.63 

338 

6.54 

4.24 

2.72 

17.09 

13.13 

14.09 

13.89 

19.09 

339 

7.05 

5.79 

3.46 

17.14 

12.35 

13.47 

13.08 

18.82 

340 

5.98 

3.93 

2.54 

16.22 

12.31 

14.12 

12.73 

17.08 

341 

6.08 

4.60 

3.46 

17.36 

12.39 

13.88 

12.26 

17.15 

342 

4.57 

2.56 

1.78 

13.59 

11.08 

11.39 

11.60 

13.63 

343 

5.56 

3.84 

2.75 

15.12 

11.12 

11.06 

11.94 

15.71 

344 

5.46 

4.37 

3.16 

• 

• 

11.98 

10.93 

14.46 

345 

4.84 

3.60 

2.50 

12.96 

9.61 

10.32 

10.15 

13.98 

346 

4.65 

3.10 

2.32 

12.45 

9.52 

10.16 

10.44 

13.06 

347 

4.48 

3.27 

2.31 

10.76 

« 

8.59 

8.34 

10.93 

348 

3.65 

2.47 

1.93 

8.44 

6.36 

7.39 

7.08 

8.65 

349 

3.31 

2.22 

1.75 

7.60 

6.08 

• 

6.70 

8.09 

350 

3.65 

2.60 

2.22 

• 

• 

• 

5.69 

7.35 

351 

3.15 

2.19 

1.78 

7.08 

5.82 

6.17 

6.12 

7.39 

352 

3.05 

2.37 

1.80 

5.81 

4.24 

• 

4.84 

6.25 

353 

2.51 

1.83 

1.37 

5.15 

3.91 

• 

4.08 

5.57 

354 

9.48 

• 

1.57 

17.53 

• 

• 

13.88 

17.34 

355 

• 

• 

• 


• 

• 


• 

356 

• 

9.08 

3.14 


• 

• 


• 

357 

• 

• 



• 

• 

• 

• 

358 

• 


• 

27.47 

20.07 

• 

21.90 

27.47 

359 

• 


3.07 

• 

• 

• 

• 

• 

360 

• 


2.97 

• 

• 

• 

• 

• 

361 

• 

• 

3.53 

27.03 

19.67 

• 

• 

27.13 

362 

8.15 

6.08 

1.91 

19.41 

14.74 

16.14 

14.24 

16.39 

363 

• 

6.36 

2.46 


13.80 

• 

13.85 

16.90 

364 

• 

• 

• 

• 

• 



• 

365 

• 

• 

• 

• 

• 



• 

366 

• 

• 

• 

• 

• 


• 

• 

367 

• 

• 

• 

• 

• 

• 

• 

• 

368 

• 

• 

• 


• 

• 

• 

• 

369 

• 

• 

• 

• 

• 

• 

• 

• 

370 

• 

• 

• 

• 

• 


• 

• 

371 

• 

9.55 

2.62 

26.75 

21.97 


21.33 

• 

372 

• 

7.92 

1.98 

• 

• 

• 

• 

• 


# 

1 

2 

3 

4 

5 

6 

7 

8 

9 

10 

11 

12 

13 

14 

15 

16 

17 

18 

19 

20 

21 

22 

23 

24 

25 

26 

27 

28 

29 

30 

31 

32 

33 

34 

35 

36 

37 

38 

39 

40 

41 

42 

43 

44 

45 

46 

47 

48 


389 


PAWI 

PAWN 

PAWI II 

PAWIV 

PAWV 

THMX 

ALI 

9.30 

9.63 

8.86 

8.55 

8.68 

7.79 

42.58 

5.43 

5.56 

5.33 

5.71 

5.39 

5.02 

23.22 

3.35 

4.16 

3.73 

4.01 

3.98 

3.91 

17.31 

3.45 

3.55 

3.83 

3.98 

3.54 

3.64 

17.31 

2.31 

2.64 

2.57 

2.65 

2.48 

3.56 

13.08 

1.95 

1.85 

2.13 

1.98 

2.04 

2.59 

10.14 

8.64 

8.17 

8.92 

8.63 

7.87 

7.09 

38.92 

4.29 

4.48 

4.62 

4.47 

4.35 

4.66 

22.94 

4.53 

4.82 

4.73 

4.59 

4.14 

4.19 

22.26 

4.64 

4.91 

4.74 

4.83 

4.77 

5.63 

22.82 

3.89 

3.94 

3.94 

4.05 

3.79 

4.80 

19.44 

3.61 

4.19 

4.24 

4.53 

3.72 

4.25 

18.55 

2.80 

2.89 

3.04 

3.00 

2.83 

4.01 

15.30 

2.69 

2.85 

2.79 

2.90 

2.71 

3.65 

• 

2.00 

2.14 

2.31 

2.19 

• 

• 

11.31 

3.40 

3.65 

4.03 

4.08 

3.42 

3.65 

18.78 

7.55 

8.63 

8.58 

8.66 

7.82 

7.74 

40.24 

7.67 

7.72 

8.00 

7.81 

7.09 

6.65 

35.05 

5.71 

6.28 

6.32 

6.13 

5.68 

6.27 

31.83 

8.14 

8.87 

• 

9.14 

8.35 

7.89 

41.19 

3.00 

3.22 

3.58 

• 

• 

3.89 

19.35 

2.70 

2.88 

• 

2.84 

• 

3.86 

14.65 

3.34 

3.60 

3.23 

3.72 

3.28 

4.02 

17.64 

3.17 

• 

3.37 

3.58 

• 

4.05 

17.60 

3.74 

4.21 

4.22 

4.20 

3.84 

3.87 

20.96 

3.06 

3.31 

3.36 

3.26 

3.04 

3.87 

• 

3.11 

3.35 

3.56 

3.59 

2.98 

3.51 

• 

4.99 

5.53 

5.67 

5.51 

5.21 

6.31 

29.46 

3.59 

3.93 

3.78 

4.21 

3.64 

4.74 

21.14 

• 

• 

• 

• 

• 

3.63 

• 

2.28 

2.61 

• 

• 

2.24 

2.83 

• 

2.36 

2.57 

2.64 

2.75 

2.41 

2.92 

• 

• 

• 

1.94 

2.01 

• 

2.13 

• 

• 

• 

• 

• 

• 

2.43 

9.29 

6.97 

7.50 

7.63 

7.77 

6.15 

6.92 

35.58 

6.36 

6.85 

7.36 

7.44 

6.38 

5.80 

31.24 

6.98 

7.02 

7.22 

7.27 

6.89 

6.61 

35.05 

2.97 

3.21 

• 

3.31 

3.30 

3.51 

17.19 

3.51 

3.40 

3.61 

• 

3.58 

3.79 

16.06 

3.36 

3.32 

3.56 

3.55 

3.23 

3.67 

16.02 

2.88 

2.89 

3.18 

3.49 

3.23 

2.99 

15.62 

2.83 

3.22 

3.17 

3.39 

• 

3.77 

14.42 

2.81 

• 

3.12 

3.07 

2.76 

3.54 

• 

2.81 

2.92 

• 

• 

2.84 

3.31 

13.01 

1.98 

• 

2.40 

2.42 

2.28 

3.02 

• 


2.74 3.04 3.13 

2.56 3.00 2.94 


2.97 

2.73 


3.74 

3.02 


ALII 

36.13 
20.40 

15.31 

14.79 

12.25 
9.66 

33.13 
20.86 
20.40 
20.18 
17.27 
16.00 
13.47 
12.57 

9.71 
17.55 

35.05 
31.52 

• 

35.90 

17.08 
12.68 
16.00 

14.63 
• 

14.22 

16.09 

27.05 
18.68 
15.72 
13.47 
10.45 

8.63 

8.71 
29.89 
28.04 

30.79 
14.99 
15.11 
14.19 

14.32 

12.35 
12.87 

12.36 

9.25 

36.37 
13.81 

11.37 


2.95 


12.29 


# 

49 

50 

51 

52 

53 

54 

55 

56 

57 

58 

59 

60 

61 

62 

63 

64 

65 

66 

67 

68 

69 

70 

71 

72 

73 

74 

75 

76 

77 

78 

79 

80 

81 

82 

83 

84 

85 

86 

87 

88 

89 

90 

91 

92 

93 

94 

95 

96 


390 


PAWI 

PAWII 

PAWI II 

PAWIV 

PAWV 

THMX 

ALI 

ALII 

1.75 

1.98 

2.12 

2.12 

1.82 

2.57 

• 

12.06 

2.31 

2.66 

2.66 

2.70 


2.81 

• 

10.84 

2.42 

2.61 

2.62 

2.61 

2.50 

2.50 

• 

10.13 

2.33 

• 

2.46 

2.47 

2.29 

2.64 

• 

10.74 

2.05 

2.31 

2.45 

• 

2.10 

2.45 

• 

• 

2.03 

2.07 


2.29 

2.19 

2.37 

• 

• 

1.70 

1.94 

2.09 

2.05 

1.70 

2.19 

• 

8.30 

1.42 

• 


1.72 

1.53 

2.26 

• 

7.79 

1.28 

1.37 

1.54 

1.46 

• 

1.79 

• 

6.21 

4.35 

4.78 

4.49 

4.96 

4.61 

5.07 

• 

19.85 

• 

• 

4.13 

4.30 

3.95 

4.29 

21.05 

19.78 

5.43 

5.52 

5.44 

5.33 

5.67 

6.22 

32.14 

27.72 

4.19 

4.07 

4.20 

4.18 

4.21 

4.69 

19.49 

17.51 

2.96 

3.18 

3.10 

3.02 

2.80 

3.72 

15.41 

14.66 

4.35 

4.45 

4.87 

4.44 

4.04 

4.68 

21.86 

20.53 

3.10 

• 

3.38 

3.49 

3.08 

4.05 

16.76 

14.46 

3.25 

3.69 

3.67 

3.74 

3.37 

4.16 

17.29 

14.82 

8.86 

8.61 

10.47 

9.17 

9.24 

8.16 

41.02 

35.66 

3.43 

3.79 

4.02 

4.24 

3.68 

3.89 

19.55 

16.51 

4.36 

4.50 

4.55 

4.44 

4.14 

5.47 

21.81 

18.93 

5.25 

4.98 

5.06 

4.89 

5.01 

6.37 

29.41 

24.53 

• 

• 


. 

• 

4.14 

16.80 

15.31 

• 

• 


• 

• 

5.15 

21.41 

17.89 

3.18 

3.27 

3.27 

3.15 

3.21 

3.45 

• 

14.54 

6.46 

6.61 

7.48 

6.64 

6.34 

6.98 

29.62 

29.67 

• 

• 

. 

• 

« 

4.46 

19.57 

16.67 

3.47 

3.77 

3.62 

3.79 

3.52 

3.50 

18.66 

17.44 

4.81 

4.95 

• 

• 

4.66 

4.75 

22.47 

19.98 

« 

• 

• 

8.93 

7.89 

• 

43.45 

37.76 

2.43 

2.83 

• 

2.70 

2.58 

3.35 

• 

12.71 

2.88 

3.28 

3.24 

• 

2.94 

3.76 

• 

13.39 

2.48 

2.82 

2.74 

2.79 

• 

3.71 

• 

11.91 

2.11 


• 

• 

• 

2.80 

13.08 

12.31 

• 

• 

• 

3.39 

2.91 

3.62 

14.68 

13.09 

• 

2.35 

2.44 

2.54 

• 

2.84 

• 

11.12 

• 


• 

• 

• 

3.62 

13.67 

12.92 

3.14 


• 

3.63 

• 

3.99 

• 

• 

• 

3.62 

3.59 

« 

2.89 

4.28 

18.56 

15.62 

2.87 

3.40 

3.20 

3.34 

3.06 

4.21 


14.17 

4.20 

4.97 

4.87 

4.92 

4.37 

5.15 

20.82 

19.68 

8.44 

8.64 

• 

8.38 

• 

• 

40.99 

36.89 

2.58 

2.66 

2.74 

2.76 

2.52 

3.46 

• 

11.88 

• 

• 

3.18 

• 

• 

3.58 

15.72 

13.58 

• 

• 

• 

• 

• 

3.05 

• 

12.75 

2.05 

2.39 

2.54 

• 

• 

2.54 

• 

12.24 

• 

3.09 

3.21 

• 

• 

3.24 

13.86 

12.06 

2.73 

3.03 

2.78 

3.14 

2.77 

3.43 

• 

10.87 

• 

2.62 

2.58 

2.55 

• 

3.08 

14.33 

12.62 


391 


# 

PAWI 

PAWN 

PAWI II 

PAWIV 

PAWV 

THMX 

ALI 

ALII 

97 

• 

• 

• 

• 

• 

3.21 

12,86 

12.01 

98 

8,77 

• 

9.79 

9.20 

• 

9.14 

42,51 

37.99 

99 

6.72 

7.35 

7.73 

7.22 

6.51 

6.62 

32.37 

28.89 

100 

8.21 

8.33 

8.47 

8.38 

7.67 

7.46 

38.15 

33.98 

101 

• 

• 

. 

• 

• 

5.69 

29.78 

26.00 

102 

6.93 

7.96 

7.91 

7.95 

7.14 

6.26 

35.21 

30.91 

103 

2.80 

2.91 

• 

3.20 

• 

3.89 

• 

13.51 

104 

2.22 

• 

• 

• 

• 

2.93 

13.05 

12.35 

105 

2,71 

2.78 

• 

• 

2.92 

3.62 

• 

11.50 

106 

• 

2.72 

• 

2,92 

• 

3.17 

• 

11.91 

107 

• 

• 

• 

• 

• 

3.44 

14.50 

13.56 

108 

• 

2.57 

• 

2.71 

• 

3.39 

14.03 

12.70 

109 

2.42 

2.77 

2.84 

2.94 

2.54 

3.11 

12.48 

11.63 

110 

• 

• 

• 

• 

3.03 

3.89 

16.49 

14.13 

111 

2.73 

• 

• 

3.04 

2.70 

3.31 

• 

12.11 

112 

1.92 

2.10 

2.03 

2.30 

2.10 

2.82 

11.15 

9.85 

113 

2.13 

2.24 

• 

2.25 

• 

2.62 

11.53 

10.10 

114 

• 

• 

• 

• 

• 

3.90 

15.25 

13.09 

115 

1.92 

2,27 

• 

• 

• 

2.88 

10.33 

9.72 

116 

2.66 

• 

• 

• 

• 

3.50 

14.31 

11.89 

117 

1.88 

2.10 

• 

2.44 

2.02 

2.51 

• 

• 

118 

. 

1.65 

1.71 

1.72 

• 

2.38 

• 

• 

119 

• 

• 

• 

• 

• 

2,51 

• 

• 

120 

7.69 

7.27 

7.92 

7.34 

7.41 

7.53 

32.80 

31.08 

121 

5.32 

5,69 

5.73 

6.33 

5.57 

5.20 

26.82 

23.22 

122 

• 

2.83 

• 

• 

2,70 

3.67 

14,22 

13.09 

123 

• 

• 

• 

2.46 

• 

2.95 

12.44 

11,56 

124 

2.19 

• 

2,59 

2.85 

2.25 

3.11 

11.79 

9.60 

125 

1.96 

2.31 

2.44 

2,59 

2.05 

2.98 

11.53 

10.23 

126 

• 

• 

• 

1.99 

• 

2.36 

9.18 

8.45 

127 

• 

• 

• 

« 

• 

• 

• 

• 

128 

3.64 

4.09 

4.51 

4.50 

3.84 

4.41 

19.20 

17.23 

129 

8,53 

8.18 

• 

• 

• 

8.21 

41,96 

37.03 

130 

• 

12.63 

11.50 

12.34 

12.48 

• 

• 

• 

131 

7.39 

7.77 

8.66 

7.87 

7.69 

7.95 

30.63 

28.32 

132 

8,02 

• 

« 

9.09 

• 

9.39 

36.55 

32.81 

133 

5,71 

6.12 

5.27 

5.66 

4.74 

8.63 

29.28 

27.44 

134 

6.10 

5.86 

5.86 

5.54 

5.54 

5.51 

• 

26.53 

135 

6.65 

6.70 

7.35 

6.79 

7.09 

8.39 

28.91 

27.35 

136 

• 

7.09 

6,43 

6.74 

6.80 

7.20 

• 

• 

137 

2,03 

2.36 

2.54 

2.51 

2.13 

2.62 

11.32 

10.38 

138 

6.79 

6.88 

7.08 

6.75 

6.88 

7.97 

36.72 

33.65 

139 

6.41 

6.52 

6.93 

6.92 

6.60 

7,48 

32.60 

30.07 

140 

5.81 

5.76 

5.86 

5.91 

5.85 

6.95 

27.96 

25.67 

141 

2.40 

2.64 

2.59 

2.89 

2,92 

2.81 

• 

• 

142 

6.78 

7.18 

7.73 

6.60 

6.71 

8.38 

• 

30.68 

143 

• 

• 

• 

• 

• 

• 

• 

• 

144 

2.61 

2.62 

2.41 

2.51 

2.15 

2.71 

11.89 

11.54 


392 


# 

PAWI 

PAWN 

PAWI II 

PAWIV 

PAWV 

THMX 

ALI 

ALII 

145 

3.42 

3.39 

3.50 

3.55 

3.30 

3.37 

16.25 

14.43 

146 

• 

• 

• 

« 

• 

• 

14.93 

13.51 

147 

8.53 

8.34 

8.83 

8.96 

9.14 

9.55 

• 

• 

148 

7.78 

8.05 

7.93 

7.84 

7.74 

8.42 

• 

• 

149 

13.09 

13.79 

13.81 

13.29 

13.41 

t 

61.95 

54.74 

150 

11.83 

10.79 

12.75 

12.12 

11.30 

• 

56.81 

50.80 

151 

11.43 

12.82 

12.85 

13.13 

11.27 

• 

52.69 

46.72 

152 

14.03 

14.70 

14.42 

14.96 

13.29 

• 

66.96 

59.25 

153 

9.84 

10.77 

9.95 

10.51 

9.99 

• 

44.92 

40.56 

154 

10.61 

11.34 

11.20 

11.60 

10.99 

• 

57.12 

50.58 

155 

• 

• 

• 

14.55 

13.90 

• 

57.72 

57.12 

156 

• 

• 

10.07 

10.20 

10.01 

• 

• 

• 

157 

10.24 

10.78 

9.95 

10.89 

10.59 

9.49 

49.94 

44.66 

158 

11.92 

12.39 

12.70 

13.10 

12.09 

• 

54.38 

50.62 

159 

9.16 

10.38 

10.28 

10.46 

9.55 

• 

45.88 

43.15 

160 

14.94 

16.04 

14.66 

17.15 

15.23 

• 

65.75 

58.05 

161 

12.89 

13.76 

13.85 

13.74 

13.11 

• 

55.06 

51.75 

162 

10.43 

11.05 

10.80 

11.23 

10.04 

• 

50.01 

45.47 

163 

9.96 

10.42 

10.54 

10.42 

9.99 

• 

45.50 

40.43 

164 

10.59 

10.64 

11.11 

10.89 

10.29 

• 

48.06 

45.30 

165 

16.01 

17.11 

17.09 

• 

16.24 

• 

64.14 

57.88 

166 

14.11 

14.95 

14.86 

15.51 

14.74 

• 

68.80 

61.01 

167 

10.55 

11.43 

10.89 

11.50 

10.93 

• 

51.68 

46.07 

168 

10.03 

11.43 

11.07 

11.09 

10.52 

9.82 

48.32 

42.54 

169 

9.58 

11.08 

11.09 

11.73 

10.83 

• 

48.75 

39.62 

170 

10.17 

12.01 

9.68 

11.66 

10.95 

• 

46.40 

• 

171 

9.94 

10.45 

10.03 

• 

9.98 

8.77 

• 

40.12 

172 

12.30 

13.13 

12.89 

13.17 

12.69 

• 

61.38 

55.41 

173 

13.16 

13.06 

13.23 

13.34 

12.25 

• 

57.87 

54.50 

174 

12.25 

12.23 

11.77 

12.38 

11.28 

• 

50.13 

45.44 

175 

8.67 

9.19 

8.99 

• 

8.80 

8.89 

44.20 

37.80 

176 

13.58 

14.67 

15.63 

15.24 

13.91 

• 

56.05 

54.14 

177 

9.82 

9.62 

9.72 

10.12 

9.53 

8.96 

• 

36.10 

178 

• 

12.11 

• 

12.30 

11.33 

• 

53.79 

• 

179 

12.59 

13.13 

13.54 

13.36 

12.44 

• 

54.63 

49.58 

180 

11.46 

11.88 

12.25 

12.50 

10.14 

• 

• 

51.61 

181 

9.53 

11.00 

10.90 

11.11 

9.74 

9.69 

46.60 

43.50 

182 

10.69 

11.04 

11.15 

11.11 

10.41 

• 

51.77 

47.34 

183 

11.78 

12.92 

11.80 

13.47 

11.98 

• 

56.29 

50.37 

184 

10.72 

11.53 

11.91 

11.66 

11.24 

• 

49.94 

43.71 

185 

12.90 

14.91 

14.80 

15.09 

12.94 

• 

59.96 

55.04 

186 

9.20 

10.17 

9.75 

10.42 

10.59 

• 

47.41 

42.21 

187 

13.44 

13.54 

13.55 

13.77 

13.37 

• 

54.17 

49.51 

188 

14.08 

15.01 

14.53 

15.06 

14.26 

• 

62.02 

55.50 

189 

12.26 

13.57 

12.99 

13.89 

12.95 

• 

55.00 

50.22 

190 

10.11 

11.56 

11.05 

12.05 

11.26 

• 

53.81 

48.21 

191 

11.15 

11.15 

11.12 

11.20 

10.96 

• 

49.98 

46.86 

192 

10.98 

11.23 

10.78 

11.22 

11.30 

• 

48.72 

43.85 


393 


# 

PAWI 

PAWN 

PAWI II 

PAWIV 

PAWV 

THMX 

ALI 

ALII 

193 

13.53 

14.49 

14.28 

15.15 

13.79 

• 

52.27 

50.66 

194 

7.22 

8.02 

7.53 

8.07 

7.41 

7.27 

35.71 

32.58 

195 

10.95 

11.96 

12.09 

12.29 

10.83 


52.75 

47.52 

196 

10.10 

11.02 

10.89 

11.06 

10.13 


45.11 

41.44 

197 

12.42 

13.99 

13.86 

14.18 

12.81 

• 

58.14 

52.01 

198 

10.19 

10.43 

10.14 

10.50 

10.03 

• 

49.68 

46.88 

199 

11.21 

11.87 

11.93 

12.40 

10.94 

• 

51.05 

46.26 

200 

11.77 

11.68 

12.17 

12.17 

11.30 

• 

51.71 

46.25 

201 

11.03 

11.74 

11.34 

12.24 

11.28 

• 

51.58 

47.31 

202 

12.54 

12.77 

13.41 

13.41 

12.81 


52.66 

51.61 

203 

8.08 

8.55 

8.78 

8.74 

8.09 

7.82 

38.34 

35.78 

204 

13.43 

14.46 

13.01 

14.70 

13.23 


67.62 

62.02 

205 

7.20 

7.91 

7.60 

8.17 

7.22 

7.23 

34.01 

33.08 

206 

15.29 

15.83 

16.73 

16.04 

15.74 

• 

69.65 

64.66 

207 

9.91 

10.14 

10.16 

10.43 

9.88 

• 

49.59 

43.99 

208 

10.62 

10.83 

10.33 

11.15 

11.02 

9.72 

• 

44.11 

209 

7.06 

7.16 

6.97 

7.15 

7.14 

6.01 

33.53 

• 

210 

7.72 

8.16 

8.02 

8.20 

7.87 

7.76 

38.74 

34.74 

211 

11.28 

12.38 

11.82 

12.76 

11.33 


50.55 

45.97 

212 

11.47 

• 

12.23 

12.69 

• 


50.12 

45.74 

213 

11.78 

13.29 

13.45 

13.43 

12.31 


52.83 

48.67 

214 

12.32 

14.54 

14.15 

14.25 

12.55 


57.47 

49.67 

215 

10.58 

12.02 


11.91 

10.94 


53.41 

47.69 

216 

13.32 

14.13 

13.53 

14.15 

14.05 

• 

59.27 

50.94 

217 

11.79 

12.21 

12.23 

12.76 

12.21 

• 

55.55 

50.50 

218 

13.36 

13.62 

13.50 

13.38 

13.23 

• 

56.86 

53.53 

219 

12.45 

13.68 

12.54 

13.30 

12.82 

• 

56.97 

50.02 

220 

11.34 

11.92 

11.26 

12.33 

11.31 

• 

47.39 

43.63 

221 

10.74 

11.07 

10.82 

11.01 

10.83 

• 

47.88 

44.76 

222 

14.09 

14.54 

15.06 

14.52 

13.85 

• 

62.82 

57.14 

223 

12.85 

13.53 

13.88 

13.58 

13.00 

• 

59.39 

51.33 

224 

17.39 

16.68 

17.85 

17.00 

17.32 

• 

• 

• 

225 

15.02 

14.16 

14.45 

14.08 

14.26 


• 

51.09 

226 

7.68 

8.16 

7.68 

7.79 

7.44 

10.24 

34.55 

31.77 

227 

21.28 

21.08 

23.11 

21.55 

21.18 

• 

88.43 

• 

228 

15.97 

14.47 

16.06 

15.24 

15.78 


74.87 

68.66 

229 

15.22 

14.42 

14.73 

14.55 

13.81 


69.39 

60.56 

230 

12.10 

12.85 

13.47 

13.13 

12.21 

12.47 

56.97 

48.60 

231 

15.18 

15.31 

15.57 

15.36 

15.18 

• 

71.79 

61.98 

232 

10.19 

10.56 

11.07 

11.12 

10.38 

11.34 

52.36 

45.68 

233 

8.08 

9.61 

9.84 

9.69 

8.72 

• 

39.19 

33.10 

234 

8.40 

8.92 

• 

9.00 

8.54 

• 

40.41 

36.09 

235 

8.55 

8.99 

9.06 

9.23 

9.11 

• 

43.29 

35.19 

236 

• 

9.25 

9.61 

9.86 

9.79 

• 

37.90 

31.38 

237 

7.73 

• 


8.35 

7.77 

9.35 

36.95 

31.88 

238 

7.63 

8.62 

8.68 

8.67 

7.96 

8.49 

31.58 

25.12 

239 

6.15 

6.78 

7.12 

6.85 

• 

7.81 

25.40 

23.20 

240 

6.69 

8.14 

8.73 

8.72 

6.92 

7.88 

31.76 

26.50 


394 


# 

PAWI 

PAWII 

241 

6.07 

6.35 

242 

5.23 

6.01 

243 

4.15 

5.11 

244 

4.77 

5.14 

245 

4.72 

4.85 

246 

4.03 

4.41 

247 

4.36 

4.74 

248 

3.87 

4.54 

249 

3.93 

• 

250 

3.45 

3.91 

251 

3.94 

4.21 

252 

• 

3.82 

253 

3.61 

4.11 

254 

• 

2.70 

255 

• 

• 

256 

3.58 

• 

257 

3.53 

3.63 

258 

3.53 

3.83 

259 

3.14 

3.53 

260 

• 

• 

261 

3.32 

3.45 

262 

2.38 

2.31 

263 

7.72 

8.19 

264 

7.73 

7.88 

265 

6.14 

6.07 

266 

10.39 

11.15 

267 

11.90 

12.50 

268 

11.60 

12.97 

269 

10.63 

11.17 

270 

• 

9.77 

271 

10.25 

11.04 

272 

7.51 

8.53 

273 

7.15 

7.23 

274 

6.40 

7.26 

275 

6.51 

7.23 

276 

4.59 

5.43 

277 

6.60 

7.80 

278 

6.48 

7.17 

279 

4.93 

5.70 

280 

7.07 

7.91 

281 

5.16 

5.66 

282 

7.49 

7.95 

283 

6.26 

6.97 

284 

• 

7.27 

285 

5.16 

5.58 

286 

5.79 

6.64 

287 

5.59 

6.34 

288 

5.10 

5.79 


PAWI II 

PAWIV 

PAWV 

5.95 

6.32 

5.89 

5.91 

6.14 

5.34 

5.14 

5.79 

5.37 

4.82 

5.34 

4.94 

4.73 

5.08 

4.90 

4.68 

4.81 

4.19 

4.64 

5.15 

4.41 

4.47 

4.95 

4.11 

4.19 

4.67 

4.24 

3.75 

3.98 

3.60 

4.08 

4.59 

4.19 

4.03 

4.57 

3.86 

2.69 

• 

• 

• 

• 

• 

3.53 

• 

3.97 

3.45 

3.63 

4.05 

3.60 

3.68 

3.73 

3.32 

3.30 

3.59 

3.16 

2.17 

2.45 

2.41 

8.39 

8.39 

7.86 

7.69 

7.56 

7.53 

5.75 

5.98 

6.20 

11.47 

11.58 

10.69 

12.70 

13.79 

10.97 

12.33 

12.80 

11.36 

10.94 

11.07 

10.69 

11.36 

11.15 

• 

8.47 

8.35 

7.69 

7.60 

7.78 

7.06 

7.57 

7.26 

6.66 

7.65 

7.06 

6.75 

5.65 

5.27 

4.49 

8.02 

8.48 

7.65 

7.34 

7.46 

6.64 

5.28 

5.94 

4.98 

7.57 

8.13 

7.56 

5.94 

5.85 

5.45 

8.09 

• 

7.41 

7.39 

6.92 

6.78 

7.10 

7.23 

6.54 

5.34 

5.61 

5.25 

7.21 

6.75 

6.26 

6.33 

6.83 

6.14 

6.19 

6.01 

5.10 


THMX 

ALI 

ALII 

7.28 

26.73 

20.09 

6.22 

24.59 

18.85 

6.31 

20.90 

16.94 

6.37 

20.26 

16.99 

6.24 

19.18 

16.51 

5.33 

18.12 

15.65 

5.81 

18.49 

15.50 

5.62 

18.65 

14.98 

4.76 

16.62 

13.39 

4.48 

16.26 

14.12 

5.37 

16.40 

13.38 

4.85 

16.38 

13.56 

4.21 

16.70 

13.30 

3.40 

11.97 

10.74 

4.78 

16.61 

14.16 

4.34 

14.68 

13.18 

4.25 

14.68 

12.73 

4.29 

14.69 

11.72 

3.82 

14.05 

11.73 

3.49 

13.22 

11.69 

3.60 

13.19 

11.22 

2.92 

10.21 

8.55 

10.15 

30.10 

26.06 

10.44 

32.61 

27.97 

7.71 

23.85 

20.11 

• 

43.99 

37.70 

• 

49.01 

42.20 

• 

• 

38.22 

• 

38.11 

36.82 

• 

38.27 

• 

10.87 

33.88 

27.19 

8.64 

31.38 

26.39 

9.77 

28.43 

23.56 

9.79 

29.03 

24.27 

5.79 

19.76 

16.21 

9.62 

29.20 

25.58 

9.17 

27.38 

22.67 

7.39 

21.33 

16.57 

8.59 

30.08 

24.51 

7.13 

25.01 

20.38 

8.02 

30.14 

24.52 

8.12 

28.09 

24.28 

8.19 

28.38 

26.00 

6.69 

21.17 

18.42 

7.27 

28.43 

23.82 

7.33 

24.16 

19.57 

6.63 

• 

19.63 


395 


# 

PAWI 

PAWN 

PAWI II 

PAWIV 

PAWV 

THMX 

ALI 

ALII 

289 

4.91 

5.17 

• 

• 

4.59 

6.55 

21.06 

18.25 

290 

6.24 

7.28 

7.05 

6.85 

6.19 

7.53 

26.92 

20.14 

291 

4.51 

5.31 

6.04 

5.55 

4.39 

6.94 

21.74 

17.64 

292 

5.10 

5.33 

5.14 

5.33 

5.41 

6.63 

19.03 

• 

293 

5.37 

6.01 

6.13 

5.96 

5.51 

6.55 

22.42 

16.76 

294 

4.59 

4.90 

5.18 

5.06 

4.79 

6.46 

19.09 

16.49 

295 

5.21 

5.83 

6.01 

5.82 

5.38 

6.64 

22.65 

19.69 

296 

5.28 

5.70 

5.71 

• 

• 

6.08 

20.37 

17.40 

297 

4.57 

4.52 

4.64 

4.88 

4.80 

6.15 

18.51 

15.12 

298 

4.50 

5.28 

5.10 

5.46 

4.91 

5.69 

19.47 

16.32 

299 

4.50 

5.34 

5.35 

5.73 

4.60 

6.47 

19.57 

16.62 

300 

3.79 

• 

« 

4.42 

3.94 

4.75 

17.44 

15.53 

301 

4.33 

4.52 

• 

4.59 

4.39 

5.25 

17.92 

15.86 

302 

3.83 

4.19 

4.42 

4.39 

4.22 

6.06 

16.82 

14.18 

303 

3.93 

4.42 

4.22 

4.48 

3.99 

5.63 

17.52 

14.20 

304 

3.59 

3.98 

3.77 

3.95 

3.82 

4.66 

15.05 

12.40 

305 

3.98 

4.20 

3.94 

4.10 

3.86 

4.47 

16.21 

12.88 

306 

3.73 

3.88 

3.81 

3.99 

3.71 

4.61 

15.28 

12.75 

307 

3.37 

3.29 

3.66 

3.49 

3.36 

3.63 

15.49 

15.31 

308 

3.49 

3.57 

3.63 

3.61 

3.32 

4.31 

16.04 

13.41 

309 

4.11 

4.49 

4.46 

4.53 

4.31 

5.78 

18.13 

15.14 

310 

3.68 

4.36 

4.22 

4.62 

3.99 

5.08 

16.44 

13.88 

311 

3.81 

4.41 

4.16 

4.15 

3.82 

4.64 

15.52 

13.57 

312 

4.11 

4.56 

4.75 

4.48 

4.42 

5.04 

17.49 

14.02 

313 

4.07 

4.79 

4.52 

5.23 

4.52 

4.73 

17.57 

15.15 

314 

5.11 

6.01 

5.88 

6.03 

5.08 

5.72 

21.10 

16.84 

315 

3.52 

3.80 

3.58 

3.80 

3.65 

4.29 

14.04 

11.96 

316 

• 

• 

• 

4.05 

3.53 

4.46 

14.81 

11.97 

317 

4.01 

4.17 

4.36 

4.50 

4.27 

5.42 

17.84 

15.11 

318 

3.44 

• 

3.61 

3.80 

3.45 

4.19 

15.21 

13.42 

319 

2.51 

2.96 

2.97 

3.03 

2.95 

3.97 

13.98 

12.09 

320 

3.08 

3.46 

3.44 

3.62 

3.28 

4.57 

13.23 

11.38 

321 

• 

• 

3.71 

3.67 

3.68 

4.82 

13.61 

11.45 

322 

• 

3.92 

4.18 

• 

3.60 

5.50 

16.61 

14.06 

323 

2.85 

3.19 

3.59 

3.79 

3.01 

3.92 

12.37 

10.48 

324 

3.47 

3.67 

3.47 

3.79 

3.55 

3.96 

13.44 

11.39 

325 

2.92 

2.89 

• 

• 

2.89 

3.69 

11.56 

9.76 

326 

2.51 

2.76 

2.77 

2.91 

2.52 

3.54 

10.65 

8.85 

327 

2.53 

2.70 

• 

2.81 

2.47 

3.13 

11.02 

9.44 

328 

2.38 

2.53 

2.51 

• 

2.49 

2.99 

10.76 

9.20 

329 

• 

2.48 

• 

2.71 

2.49 

3.38 

10.21 

8.71 

330 

2.11 

• 

• 

2.34 

2.14 

2.89 

9.64 

8.26 

331 

2.18 

2.34 

• 

2.34 

2.18 

3.05 

9.74 

8.58 

332 

• 

• 

• 

2.26 

2.24 

2.80 

9.93 

8.48 

333 

1.96 

1.93 

• 

2.11 

2.14 

2.92 

9.21 

8.17 

334 

9.64 

10.79 

10.97 

11.01 

9.57 

12.67 

44.90 

37.02 

335 

7.10 

7.84 

• 

8.22 

7.07 

8.98 

29.41 

25.42 

336 

6.02 

6.01 

6.83 

6.68 

6.57 

9.58 

26.28 

21.79 


396 


# 

PAWI 

PAWN 

PAWI II 

PAWIV 

337 

5.68 

6.04 

6.38 

6.53 

338 

5.03 

5.90 

6.02 

6.16 

339 

5.25 

5.53 

5.61 

5.48 

340 

4.97 

5.38 

5.12 

5.70 

341 

5.20 

5.64 

6.16 

5.84 

342 

4.32 

4.05 

4.48 

4.36 

343 

4.48 

4.89 

4.77 

4.89 

344 

• 

« 

4.57 

5.17 

345 

4.03 

4.43 

4.30 

4.65 

346 

4.11 

4.87 

4.48 

4.92 

347 

3.29 

• 

3.96 

4.12 

348 

2.75 

3.07 

2.72 

2.96 

349 

2.47 

2.60 


2.79 

350 

• 

• 


3.06 

351 

2.44 

2.57 

2.48 

2.58 

352 

2.03 

2.10 


2.27 

353 

1.57 

1.83 


1.98 

354 

8.29 

7.84 

• 

8.58 

355 

• 

• 

• 

• 

356 


10.85 

11.63 

• 

357 


• 

• 

• 

358 

11.21 

11.12 

10.83 

11.67 

359 

* 

• 

• 

• 

360 

• 

• 

• 

• 

361 

10.73 

11.23 

10.96 

10.60 

362 

7.50 

6.69 

7.15 

7.55 

363 

6.09 

6.50 

6.87 

6.99 

364 

• 

• 

• 


365 

• 


• 


366 

• 

• 

• 


367 

• 

• 

• 


368 

• 

• 

• 


369 

• 

• 

• 


370 

• 

• 

• 


371 

12.78 

11.84 

• 

12.86 

372 

• 

• 

• 

• 


PAWV 

THMX 

ALI 

ALII 

5.91 

8.39 

24.84 

20.31 

5.67 

6.89 

23.02 

18.96 

5.37 

8.51 

22.87 

18.05 

5.26 

6.82 

21.62 

17.97 

5.57 

6.79 

21.64 

17.40 

4.71 

4.75 

19.74 

19.64 

4.57 

6.17 

19.18 

16.70 

4.42 

6.32 

19.17 

15.08 

4.31 

5.05 

17.21 

15.46 

4.38 

4.95 

16.66 

13.99 

3.34 

4.88 

14.65 

12.22 

3.01 

4.07 

12.25 

10.58 

2.72 

3.38 

11.95 

10.28 

3.19 

3.82 

11.89 

10.83 

2.65 

3.42 

10.82 

9.71 

2.30 

3.35 

9.78 

8.53 

1.68 

2.62 

7.47 

6.17 

8.17 

10.68 

33.05 

30.14 

• 

• 

• 

• 

44.71 

40.29 

• 

• 

11.30 

• 

• 

• 

• 

• 

• 

• 

10.90 

• 

• 

• 

• 

• 

8.12 

8.07 

33.64 

30.83 

7.23 

• 

• 

• 

• 

• 

35.77 

28.02 

• 

• 

14.56 

11.40 

• 

• 

13.50 

10.23 

• 

• 

12.82 

9.95 

• 

• 

12.11 

9.94 


77.96 


67.52 


397 


# 

ALIN 

ALIV 

ALV 

IL1 

IL2 

IL3 

IL4 

IL5 

1 

32.13 

35.20 

42.08 

38.81 

34.04 

32.79 

38.45 

40.56 

2 

19.72 

20.47 

23.34 

22.73 

20.35 

20.52 

22.84 

22.93 

3 

14.13 

15.10 

17.46 

16.76 

14.85 

14.37 

16.61 

17.71 

4 

14.10 

15.20 

17.60 

16.57 

13.93 

13.94 

16.04 

17.46 

5 

10.98 

12.05 

13.01 

13.58 

11.77 

11.48 

12.97 

12.86 

6 

9.29 

9.79 

9.96 

11.01 

9.48 

9.41 

10.85 

10.98 

7 

31.86 

32.60 

38.06 

38.29 

32.89 

32.04 

36.29 

37.84 

8 

18.50 

20.56 

22.94 

23.05 

19.73 

18.79 

22.72 

23.20 

9 

18.96 

19.16 

21.85 

21.45 

19.34 

19.16 

20.70 

21.60 

10 

18.40 

19.13 

22.72 

23.05 

19.90 

18.89 

22.07 

22.49 

11 

15.54 

17.24 

19.71 

19.72 

16.97 

16.51 

19.92 

20.06 

12 

14.13 

16.14 

19.24 

16.99 

14.74 

14.80 

17.33 

18.58 

13 

12.56 

13.41 

15.36 

15.70 

13.10 

12.62 

14.88 

15.72 

14 

11.68 

12.67 

• 

14.14 

12.75 

12.25 

13.88 

14.41 

15 

8.45 

9.49 

10.90 

10.70 

8.72 

8.35 

9.94 

10.88 

16 

• 

• 

• 

19.49 

16.70 

• 

19.03 

19.77 

17 

32.41 

34.68 

40.10 

37.13 

32.29 

32.16 

35.92 

38.74 

18 

29.07 

30.73 

34.97 

32.18 

28.54 

28.25 

31.80 

33.12 

19 

26.64 

28.23 

32.38 

31.59 

27.58 

26.78 

• 

29.54 

20 

32.71 

35.24 

41.16 

37.80 

32.80 

32.02 

36.77 

38.13 

21 

15.13 

16.89 

19.49 

19.38 

15.96 

15.92 

18.99 

19.40 

22 

11.77 

12.25 

14.38 

13.71 

11.55 

11.78 

13.74 

13.98 

23 

15.41 

15.69 

17.88 

17.37 

15.95 

15.79 

16.64 

17.94 

24 

13.53 

14.96 

17.72 

17.62 

14.35 

13.83 

17.09 

18.00 

25 

17.71 

18.64 

21.13 

• 

18.59 

17.60 

20.11 

21.62 

26 

13.29 

14.24 

• 

16.38 

14.27 

14.13 

• 

16.33 

27 

14.13 

15.22 

• 

17.20 

15.50 

14.70 

16.53 

• 

28 

25.22 

25.92 

28.95 

29.07 

25.84 

25.10 

27.63 

27.40 

29 

17.01 

17.94 

20.85 

20.72 

17.51 

17.17 

19.36 

20.70 

30 

14.24 

15.41 

• 

17.72 

15.74 

14.93 

• 

18.13 

31 

12.25 

13.38 

• 

14.96 

12.85 

12.39 

14.63 

14.98 

32 

9.91 

10.42 

• 

11.73 

10.51 

10.14 

11.36 

11.49 

33 

8.33 

8.64 

• 

9.65 

8.67 

8.25 

9.62 

10.18 

34 

7.87 

8.10 

9.56 

9.28 

8.29 

8.10 

9.18 

9.80 

35 

27.60 

28.96 

35.86 

33.03 

27.99 

27.77 

31.73 

34.39 

36 

25.52 

27.36 

31.41 

28.86 

25.61 

25.05 

27.96 

30.08 

37 

28.52 

30.25 

35.03 

33.80 

29.09 

28.06 

33.12 

33.90 

38 

13.91 

16.28 

17.43 

17.42 

14.47 

14.79 

17.53 

18.04 

39 

13.36 

14.56 

16.25 

16.14 

14.52 

14.04 

15.87 

16.01 

40 

12.77 

13.56 

15.68 

15.18 

13.39 

13.09 

14.36 

15.63 

41 

13.10 

14.03 

16.15 

15.11 

13.34 

12.94 

14.54 

16.33 

42 

11.82 

12.29 

14.83 

14.30 

12.87 

12.75 

14.65 

14.71 

43 

11.79 

12.89 

• 

13.99 

11.96 

11.97 

14.21 

15.31 

44 

11.44 

11.93 

13.69 

12.91 

11.82 

11.50 

13.42 

13.43 

45 

8.25 

8.90 

• 

10.68 

8.99 

8.96 

10.50 

11.39 

46 

35.06 

34.67 

42.04 

39.64 

34.69 

34.54 

37.74 

• 

47 

• 

13.81 

• 

14.96 

13.25 

• 

15.03 

15.62 

48 

10.55 

11.22 

12.14 

12.49 

10.75 

10.18 

12.19 

13.18 


398 


# 

ALIII 

ALIV 

ALV 

IL1 

IL2 

IL3 

IL4 

IL5 

49 

10.74 

11.64 

• 

12.25 

11.55 

10.90 

12.82 

12.90 

50 

10.38 

11.43 

• 

12.89 

9.77 

10.83 

12.64 

13.38 

51 

9.56 

9.99 

• 

11.31 

9.58 

9.43 

11.32 

12.12 

52 

9.20 

10.33 

10.97 

11.89 

9.69 

9.77 

11.20 

11.50 

53 

9.06 

• 

• 

9.20 

8.68 

9.05 

9.42 

8.80 

54 

8.59 

• 

• 

10.32 

8.91 

8.92 

10.05 

9.85 

55 

7.97 

8.24 

• 

9.19 

8.14 

8.02 

9.01 

9.60 

56 

7.08 

• 

• 

8.61 

7.67 

6.99 

8.34 

8.58 

57 

6.32 

6.65 

• 

6.93 

6.42 

6.50 

7.21 

7.20 

58 

19.05 

20.07 

22.64 

21.72 

19.39 

19.10 

20.67 

22.32 

59 

18.00 

19.63 

21.19 

21.01 

18.88 

18.19 

20.65 

21.31 

60 

25.65 

28.18 

• 

28.87 

24.66 

25.28 

29.12 

29.83 

61 

16.86 

17.04 

19.83 

19.71 

17.38 

17.09 

19.15 

19.81 

62 

14.22 

14.47 

15.87 

16.25 

14.79 

14.70 

16.06 

16.40 

63 

18.11 

19.43 

21.97 

23.08 

19.43 

19.68 

22.26 

23.21 

64 

13.93 

14.70 

16.72 

15.97 

14.22 

14.04 

16.19 

16.91 

65 

14.75 

15.01 

17.09 

17.09 

14.96 

14.98 

17.74 

17.38 

66 

33.68 

34.86 

42.36 

38.65 

33.28 

33.57 

37.98 

39.67 

67 

15.17 

16.33 

19.93 

17.75 

• 

15.36 

17.95 

19.97 

68 

18.03 

19.10 

22.27 

20.94 

18.79 

18.46 

21.23 

21.78 

69 

22.65 

23.93 

28.68 

27.13 

23.60 

22.63 

26.73 

27.68 

70 

• 

14.78 

• 

17.39 

15.01 

14.32 

17.13 

• 

71 

17.22 

18.05 

21.71 

19.64 

17.37 

16.77 

20.19 

21.03 

72 

13.81 

14.88 

• 

16.31 

14.47 

14.40 

16.56 

17.55 

73 

28.70 

29.33 

• 

30.68 

28.80 

28.98 

30.14 

• 

74 

15.40 

16.67 

• 

18.49 

16.53 

16.09 

18.64 

18.97 

75 

15.54 

16.94 

18.24 

19.32 

16.42 

15.81 

18.65 

18.99 

76 

18.77 

18.71 

21.66 

20.04 

19.01 

19.05 

21.20 

22.28 

77 

35.61 

37.79 

44.06 

39.68 

35.10 

36.14 

40.08 

40.39 

78 

12.11 

12.50 

• 

13.80 

12.76 

12.56 

13.84 

14.17 

79 

12.66 

13.15 

• 

15.35 

13.34 

13.08 

15.01 

15.79 

80 

11.20 

11.91 

• 

13.75 

12.16 

11.98 

13.70 

13.62 

81 

11.34 

11.76 

12.97 

13.34 

11.98 

11.48 

13.33 

13.88 

82 

11.95 

12.49 

14.36 

13.69 

12.53 

11.93 

13.41 

15.06 

83 

10.54 

11.22 

• 

12.25 

10.69 

10.82 

12.49 

12.06 

84 

12.31 

12.92 

14.19 

14.22 

12.92 

12.57 

14.45 

14.42 

85 

19.13 

• 

• 

• 

19.48 

19.05 

• 

21.94 

86 

14.28 

16.00 

• 

17.11 

15.03 

15.12 

• 

19.06 

87 

12.70 

14.12 

• 

15.49 

13.51 

13.55 

15.79 

15.91 

88 

18.03 

19.41 

22.14 

21.22 

18.87 

18.51 

20.76 

21.71 

89 

34.31 

36.25 

• 

40.74 

34.27 

33.46 

39.06 

41.92 

90 

10.80 

• 

• 

13.60 

11.35 

11.26 

• 

13.00 

91 

12.70 

13.33 

15.45 

14.89 

12.96 

13.06 

14.69 

14.64 

92 

11.82 

12.42 

• 

13.97 

12.66 

12.59 

13.97 

13.95 

93 

11.16 

12.19 

• 

13.25 

11.69 

11.79 

13.10 

12.72 

94 

11.43 

12.05 

14.26 

13.30 

11.88 

11.65 

13.60 

13.62 

95 

11.07 

10.65 

• 

13.23 

11.41 

11.22 

13.11 

14.05 

96 

12.05 

12.85 

13.85 

13.89 

12.56 

12.44 

13.81 

14.83 


399 


# 

ALIII 

ALIV 

ALV 

IL 1 

IL 2 

IL 3 

IL 4 

IL 5 

97 

11.16 

11.65 

13.05 

13.13 

11.77 

11.68 

13.36 

13.13 

98 

36.34 

36.83 

42.60 

39.87 

36.09 

34.64 

39.14 

40.70 

99 

25.86 

. 

32.18 

29.84 

25.76 

25.52 

29.07 

31.17 

100 

30.53 

32.46 

37.84 

37.19 

31.99 

31.19 

34.89 

36.70 

101 

23.90 

25.81 

29.70 

27.92 

23.99 

23.90 

27.58 

28.39 

102 

29.10 

30.14 

34.08 

31.41 

28.85 

28.28 

31.66 

32.84 

103 

12.56 

13.53 

• 

14.98 

13.51 

13.39 

14.98 

15.34 

104 

11.76 

11.89 

• 

13.10 

11.87 

11.32 

• 

14.27 

105 

10.64 

11.79 

t 

13.08 

11.68 

11.73 

13.10 

13.18 

106 

10.76 

12.09 

t 

13.34 

11.58 

11.25 

13.50 

14.18 

107 

12.10 

12.64 

14.41 

15.34 

12.91 

12.45 

14.66 

14.68 

108 

10.88 

11.82 

13.99 

13.97 

• 

11.07 

14.03 

13.61 

109 

10.59 

11.63 

12.85 

12.95 

11.04 

11.02 

12.97 

13.72 

110 

13.33 

14.30 

16.00 

15.49 

13.37 

13.56 

15.26 

16.20 

111 

11.39 

11.72 

. 

13.62 

11.73 

11.68 

13.62 

13.52 

112 

9.75 

9.81 

11.01 

10.82 

10.00 

9.62 

10.85 

11.21 

113 

9.55 

• 

• 

11.39 

9.70 

9.49 

11.32 

11.83 

114 

12.21 

13.06 

15.31 

14.68 

12.82 

12.43 

14.75 

15.57 

115 

8.99 

9.82 

10.59 

10.43 

9.65 

9.22 

10.62 

11.21 

116 

11.93 

11.93 

14.36 

14.21 

12.64 

12.44 

14.37 

14.32 

117 

9.02 

• 

• 

10.16 

9.65 

9.00 

9.95 

10.52 

118 

• 

• 

• 

• 

• 

• 

• 

• 

119 

• 

• 

• 

• 

• 

• 

• 

• 

120 

27.10 

30.94 

34.45 

31.86 

27.71 

27.38 

32.32 

33.08 

121 

20.92 

22.39 

26.78 

24.54 

21.79 

20.81 

24.40 

25.47 

122 

. 

12.72 

14.21 

14.02 

12.16 

12.11 

13.75 

14.41 

123 

10.38 

11.36 

12.36 

12.44 

10.50 

10.49 

12.20 

12.80 

124 

9.27 

10.08 

12.11 

11.32 

9.90 

9.82 

11.83 

12.19 

125 

9.93 

10.26 

11.65 

11.58 

10.16 

9.90 

11.32 

12.38 

126 

8.15 

8.35 

9.22 

9.60 

8.30 

8.20 

9.39 

10.10 

127 

36.04 

39.17 

• 

• 

36.84 

36.99 

42.36 

• 

128 

15.30 

17.76 

19.32 

18.59 

16.20 

15.91 

18.91 

19.67 

129 

34.23 

35.85 

43.07 

39.11 

34.54 

33.82 

38.45 

39.99 

130 

• 

• 

• 

• 

• 

• 

• 

• 

131 

26.70 

26.54 

31.67 

30.30 

28.28 

27.03 

30.63 

31.52 

132 

32.46 

32.75 

• 

34.81 

32.10 

31.43 

35.39 

35.29 

133 

26.02 

• 

29.70 

27.73 

26.75 

25.19 

27.62 

28.85 

134 

25.37 

28.00 

• 

• 

25.42 

25.79 

27.87 

• 

135 

27.40 

26.28 

• 

28.82 

27.73 

26.58 

28.63 

29.26 

136 

• 

• 

• 

• 

• 

• 

• 

• 

137 

9.77 

10.47 

11.17 

11.30 

9.96 

9.88 

11.27 

11.48 

138 

31.19 

31.95 

36.27 

34.63 

31.39 

30.32 

34.01 

34.92 

139 

29.26 

29.42 

32.72 

32.89 

29.36 

29.05 

31.91 

32.66 

140 

24.30 

25.47 

28.23 

27.91 

26.07 

25.78 

28.01 

28.24 

141 

• 

• 

• 

• 

• 

• 

• 

• 

142 

30.23 

29.45 

34.13 

33.43 

30.23 

28.95 

33.64 

32.82 

143 

• 

• 

• 

• 

• 

• 

• 

• 

144 

10.94 

11.46 

12.36 

12.25 

11.26 

10.93 

12.26 

13.23 


400 


# 

ALIII 

ALIV 

ALV 

IL1 

IL2 

IL3 

IL4 

IL5 

145 

13.37 

14.17 

16.44 

15.69 

13.81 

13.61 

15.50 

16.17 

146 

12.64 

13.58 

14.64 

14.74 

13.10 

12.81 

14.71 

14,26 

147 

• 

• 


• 

• 

« 

• 

• 

148 

• 

• 

• 

• 

• 

• 

• 

• 

149 

48.42 

54.11 

• 

61.27 

51.33 

51.07 

56.68 

55.00 

150 

46.67 

51.37 

56.24 

51.06 

48,57 

48.95 

52.80 

50.87 

151 

42.31 

46.44 

52,55 

52.13 

43.92 

43.82 

49.87 

51,11 

152 

53.28 

59.15 

64.86 

59.01 

55.35 

54.90 

58.76 

61.27 

153 

35.46 

39.20 

43.80 

42.46 

37.94 

36.53 

40.64 

43.64 

154 

46.02 

51.07 

55.98 

50.86 

46.55 

47.02 

51.54 

53.50 

155 

53.30 

57.53 

62.31 

60.15 

53.95 

54,71 

61.03 

59.21 

156 

33.35 

36.54 

41.07 

• 

35,59 

35.42 

37.67 

• 

157 

42.78 

43.81 

51.04 

• 

43.71 

43.04 

45.14 

49.39 

158 

45.96 

49.82 

54.78 

53.18 

51.06 

49.45 

53.08 

52.89 

159 

38.77 

42.32 

45.09 

44,60 

40.71 

40.09 

44.06 

41.77 

160 

53.71 

58.02 

64,28 

59.16 

53.81 

54.60 

58.45 

57.98 

161 

48.76 

52.37 

57.34 

56.73 

51.30 

49.39 

56.83 

53.86 

162 

42.27 

45.78 

50.25 

48.53 

44.60 

43.87 

47.75 

48.69 

163 

37.03 

39.76 

45.17 

43,73 

37.74 

38.42 

42.75 

43.33 

164 

41.36 

45.30 

50,12 

44.28 

43.04 

41,79 

46.65 

46.36 

165 

50.09 

54.89 

60.97 

59.08 

56.79 

52.85 

55.26 

60.19 

166 

54,84 

• 

« 

61.73 

58.45 

• 

• 

65.61 

167 

43,00 

46.18 

52.38 

48.04 

44,76 

44.99 

47.73 

49.82 

168 

38.54 

41.65 

46.76 

46.18 

39.80 

39,19 

44.46 

45.25 

169 

41.79 

45.85 

49.06 

47.07 

39.71 

46.30 

46.97 

45.89 

170 

39.90 

42.20 

45.88 

• 

• 

42.27 

44.08 

• 

171 

36.25 

39.42 

42.96 

40.60 

39.51 

38.22 

41.40 

• 

172 

48.84 

54.25 

60.92 

60.94 

55.23 

53.79 

59.86 

58.84 

173 

48.37 

52.59 

56.40 

54.44 

51.76 

49.16 

55.13 

51.23 

174 

41.63 

42.96 

49.58 

46.69 

42.97 

41.40 

45.31 

• 

175 

35.40 

37.56 

42.58 

40.24 

37.08 

36.93 

39.76 

41.00 

176 

49.71 

52,80 

57.01 

59.24 

52.78 

51.14 

57.97 

50.95 

177 

32.92 

35,43 

40.46 

38.47 

35,48 

34.31 

37.80 

38.03 

178 

44.41 

46.42 

52.38 

• 

45.28 

43.89 

• 

48.96 

179 

43.23 

49.79 

55.90 

49.73 

46.61 

46.67 

51.05 

50.33 

180 

46.07 

52.32 

57.79 

53.80 

50,87 

49.49 

54.67 

53.96 

181 

40.60 

• 

45.64 

43.91 

43.05 

40.79 

42.33 

42.98 

182 

43.12 

45.61 

50.43 

49.27 

46.75 

44.08 

47.02 

49.30 

183 

45.37 

49.90 

56.75 

51.77 

47.30 

46.40 

51.93 

52.49 

184 

41.12 

46.11 

49.87 

46.78 

42.94 

43.16 

47.52 

46,34 

185 

53.18 

55.14 

60.19 

58.19 

56.49 

55.48 

58.60 

54.12 

186 

40,13 

42.37 

46.60 

43.96 

37.81 

40.55 

43.55 

44,61 

187 

47.25 

49.25 

55.31 

50.69 

48.35 

46.49 

52.45 

50.17 

188 

52.50 

54.36 

62.07 

57.55 

52.17 

52.53 

57.71 

56.56 

189 

46.78 

50.12 

53.70 

52.23 

47.96 

48.00 

52.05 

49,44 

190 

41.98 

47.13 

52.87 

52.65 

45.24 

45.69 

50.03 

50.30 

191 

41.44 

46.29 

51.87 

49.31 

46.71 

44.84 

50.33 

48.90 

192 

41.08 

43.16 

48.72 

46.17 

41.36 

41.19 

45.78 

43.98 


401 


# 

ALIII 

ALIV 

ALV 

IL1 

IL2 

IL3 

IL4 

IL5 

193 

47.68 

51.06 

52.37 

49.61 

48.50 

49.14 

50.92 

48.50 

194 

30,48 

32.43 

35.28 

34.29 

32.84 

32.20 

34.53 

33.78 

195 

43.03 

46.30 

52.67 

51.08 

48.33 

47.86 

52.41 

52.11 

196 

37.32 

40.64 

44.80 

42.03 

39.63 

38.84 

42.02 

41.30 

197 

46.85 

50.64 

57.12 

54.16 

49.98 

49.65 

53.34 

54.91 

198 

39.00 

45.16 

48.74 

48.27 

43.21 

42.07 

47.04 

44.94 

199 

41.55 

46.48 

51.58 

46.83 

43.40 

42.95 

47.25 

48.46 

200 

41.96 

46.86 

51.34 

49.19 

44.47 

44.79 

49.97 

46.95 

201 

42.67 

45.10 

50.57 

47.58 

45.04 

43.88 

46.36 

47,90 

202 

46,07 

50.48 

52.93 

51.34 

47.82 

46.93 

51.71 

46.57 

203 

33.51 

35.02 

39.33 

36.46 

32.96 

33.19 

36.46 

35.98 

204 

52.37 

60.57 

69.41 

59.61 

54.45 

52.93 

61.09 

62.58 

205 

31.62 

31.91 

35.31 

34.82 

32.53 

32.44 

34.44 

32.89 

206 

59.43 

64,83 

70.94 

66.71 

61.29 

60.71 

67.14 

66.85 

207 

41.38 

45.44 

49.89 

45.38 

44.22 

42.04 

47.71 

46.97 

208 

37.87 

43.20 

51.59 

45.64 

40.16 

39.08 

44.82 

• 

209 

27.20 

30.92 

32.97 

32.73 

28.99 

28.89 

31.97 

31.63 

210 

31.68 

32,82 

39.84 

36.59 

33.23 

32.47 

37.26 

37,51 

211 

41.35 

45.44 

50.80 

48.22 

45.52 

44.72 

48.24 

48.76 

212 

41.72 

44,35 

51.08 

47,17 

43.91 

42.42 

46.54 

47.16 

213 

44.93 

48.13 

51.79 

53.43 

49.26 

47.75 

52.67 

52.58 

214 

45.90 

49.01 

56.23 

53.36 

48.76 

47,42 

53.74 

53.33 

215 

44.01 

46.36 

52.27 

47.86 

45.70 

45.41 

48.10 

47.21 

216 

45.70 

51.05 

• 

54.59 

44.09 

46.84 

53.91 

• 

217 

42.44 

49.22 

54.94 

52.59 

46.09 

46.12 

51.62 

• 

218 

47.03 

52.34 

56.78 

51,94 

49.11 

47.86 

50.68 

49.25 

219 

44.53 

49.44 

55.00 

53.54 

44,98 

46.25 

51.48 

53.03 

220 

39.99 

43.31 

45.72 

46.16 

43.18 

41.79 

45.61 

47.28 

221 

40.04 

44.01 

47.69 

47.13 

42.73 

41.84 

46.77 

47.32 

222 

47.93 

56.25 

63.21 

58.48 

51.50 

49.95 

• 

58.78 

223 

43.33 

51.62 

60.70 

53.31 

46,01 

45.49 

51.76 

• 

224 

• 

• 

• 

• 

• 

• 

• 

• 

225 

43,44 

51.32 

• 

55.52 

45.98 

45.55 

53.63 

• 

226 

28.35 

31,87 

35.22 

35.82 

31.44 

30.74 

• 

32.55 

227 

69.04 

79.97 

86.58 

78.06 

72.55 

71.23 

79.64 

77.54 

228 

56.61 

68.54 

75.61 

66.70 

62.83 

60.49 

65.92 

67.17 

229 

51.33 

59.20 

69.71 

65.21 

57.22 

56.61 

64.57 

63.77 

230 

42.77 

47.98 

57.26 

55,46 

46,27 

45.47 

53,47 

53.98 

231 

50.88 

61.62 

71.46 

66.52 

56.29 

55.33 

65.55 

• 

232 

40.29 

46.85 

53.21 

46.76 

43.04 

42.95 

47.89 

49.27 

233 

38.97 

32.63 

38.86 

45.02 

42.24 

41.61 

44.99 

47.72 

234 

39,86 

34.04 

40.17 

47.42 

45.23 

42.22 

47.63 

47,15 

235 

38.78 

34.98 

42.35 

48.90 

42.43 

41.74 

45.32 

51.37 

236 

34.88 

29.75 

38.70 

42.64 

38.31 

36.69 

41.77 

47.71 

237 

35.54 

30.56 

35.59 

43.29 

39,48 

38.36 

39.82 

43.92 

238 

28.18 

24.47 

32.04 

34.36 

31.54 

30.74 

33.89 

38.55 

239 

24.23 

22.77 

26.14 

28.53 

26.09 

26.20 

28.46 

29.61 

240 

30.86 

27.40 

31.82 

37.79 

34.23 

34.37 

36.93 

38.87 


402 


# 

ALIII 

ALIV 

ALV 

111 

IL 2 

IL 3 

IL 4 

IL 5 

241 

24.51 

20.80 

25.42 

30.27 

27.10 

27.30 

28.80 

31.54 

242 

20.26 

18.35 

24.56 

26.96 

22.12 

21.61 

24.68 

30.73 

243 

19.82 

• 

20.47 

23.76 

20.20 

20.70 

21.79 

25.08 

244 

17.98 

16.15 

19.73 

22.27 

19.87 

19.25 

20.78 

23.86 

245 

17.60 

16.42 

19.25 

20.95 

19.17 

18.75 

19.59 

22.44 

246 

16.70 

15.35 

18.49 

20.14 

17.36 

17.75 

19.51 

23.66 

247 

16.35 

15.03 

18.00 

19.43 

17.47 

16.96 

18.40 

21.38 

248 

16.45 

14.47 

18.41 

19.15 

17.69 

17.20 

17.98 

21.38 

249 

15.62 

13.18 

16.58 

18.70 

16.73 

16.29 

17.80 

21.17 

250 

14.73 

13.70 

16.57 

19.58 

16.61 

16.47 

18.40 

21.03 

251 

15.82 

13.47 

16.20 

18.82 

16.89 

16.49 

17.42 

20.34 

252 

14.36 

13.22 

16.78 

16.47 

15.50 

15.17 

16.52 

19.31 

253 

14.46 

12.94 

16.34 

17.86 

15.39 

15.82 

17.00 

19.76 

254 

10.36 

10.41 

12.21 

12.35 

10.74 

10.69 

11.62 

13.42 

255 

14.04 

13.72 

16.56 

17.44 

15.35 

15.20 

16.56 

19.24 

256 

13.15 

12.42 

14.59 

15.87 

13.72 

13.52 

14.49 

16.68 

257 

13.72 

12.24 

14.28 

15.70 

14.26 

■ 14.47 

14.68 

16.71 

258 

13.10 

11.54 

14.82 

14.98 

13.46 

13.70 

14.31 

17.83 

259 

11.78 

11.50 

13.99 

14.73 

13.01 

12.61 

13.94 

15.91 

260 

12.49 

11.26 

13.22 

14.08 

12.91 

12.57 

13.56 

15.34 

261 

11.83 

10.69 

12.80 

13.89 

11.96 

11.93 

12.54 

15.35 

262 

8.92 

8.10 

9.93 

10.55 

9.04 

8.91 

9.57 

11.22 

263 

27.49 

25.13 

30.82 

36.26 

32.08 

31.17 

34.71 

37.41 

264 

29.66 

27.95 

32.50 

36.11 

32.92 

32.22 

36.47 

39.02 

265 

22.71 

19.57 

23.75 

26.55 

23.81 

23.32 

25.80 

28.78 

266 

• 

39.28 

43.94 

• 

• 

• 

50.31 

52.42 

267 

42.31 

37.05 

44.71 

49.75 

47.75 

46.94 

50.81 

54.78 

268 

46.69 

42.87 

48.95 

53.67 

52.12 

51.23 

56.03 

58.25 

269 

41.43 

37.71 

46.25 

49.27 

47.76 

46.05 

51.77 

55.81 

270 

39.19 

36.06 

42.92 

50.02 

46.31 

45.59 

51.77 

51.92 

271 

• 

36.24 

• 

45.80 

• 

41.54 

• 

• 

272 

31.33 

26.16 

34.10 

37.98 

34.41 

33.70 

37.09 

41.39 

273 

28.90 

25.14 

30.39 

33.64 

30.70 

29.81 

32.50 

38.02 

274 

25.03 

24.23 

30.05 

32.13 

28.15 

28.60 

33.04 

34.27 

275 

27.53 

24.52 

29.28 

32.38 

30.22 

30.21 

33.79 

35.07 

276 

17.15 

• 

18.73 

21.66 

19.04 

18.70 

21.21 

23.65 

277 

29.18 

25.85 

29.43 

34.32 

30.77 

31.00 

34.67 

35.06 

278 

25.31 

23.43 

28.24 

30.01 

27.30 

27.79 

30.97 

33.56 

279 

19.32 

16.58 

21.21 

22.79 

20.52 

19.24 

22.54 

27.31 

280 

26.98 

24.39 

29.33 

33.16 

29.71 

29.39 

33.97 

38.16 

281 

22.19 

19.75 

24.96 

27.76 

24.59 

23.99 

27.02 

29.64 

282 

27.44 

23.46 

29.92 

34.29 

30.28 

29.44 

33.54 

37.05 

283 

27.11 

24.56 

27.84 

33.29 

29.39 

29.46 

32.81 

35.70 

284 

26.86 

24.17 

28.07 

31.41 

29.98 

28.58 

30.35 

33.12 

285 

20.41 

17.55 

21.46 

22.94 

• 

21.15 

• 

25.10 

286 

24.78 

23.89 

28.59 

32.01 

28.70 

28.58 

33.04 

34.65 

287 

23.01 

19.55 

23.94 

26.62 

24.55 

24.53 

25.63 

30.77 

288 

21.03 

18.61 

22.39 

• 

22.50 

22.09 

24.31 

• 


403 


# 

ALIM 

ALIV 

ALV 

IL1 

IL2 

IL3 

IL4 

IL5 

289 

19.49 

17.79 

20.86 

23.88 

21.10 

20.45 

22.83 

26.45 

290 

21.80 

20.08 

25.74 

29.85 

25.60 

25.22 

29.58 

31.08 

291 

20.06 

17.20 

21.40 

24.45 

21.87 

21.08 

24.93 

26.28 

292 

• 

• 

• 

• 

• 

• 

• 

• 

293 

18.07 

17.12 

22.48 

23.66 

19.82 

19.74 

22.45 

27.13 

294 

18.15 

16.27 

19.54 

20.69 

19.29 

18.73 

20.87 

23.64 

295 

21.08 

19.03 

22.77 

25.99 

23.57 

23.26 

24.72 

27.62 

296 

19.32 

17.20 

20.54 

21.57 

20.58 

20.34 

22.02 

23.03 

297 

16.71 

15.52 

19.27 

19.27 

17.26 

17.65 

20.11 

22.31 

298 

18.26 

15.76 

19.75 

20.83 

20.00 

19.50 

19.99 

23.02 

299 

18.60 

15.56 

20.24 

22.18 

20.03 

19.56 

21.64 

24.20 

300 

15.88 

15.15 

17.80 

19.27 

17.17 

16.76 

18.83 

19.95 

301 

16.33 

15.51 

17.97 

19.40 

17.75 

17.09 

19.20 

20.75 

302 

15.00 

13.82 

17.04 

18.24 

16.49 

15.97 

17.88 

19.28 

303 

15.92 

14.33 

17.50 

19.13 

17.15 

16.32 

18.59 

20.63 

304 

13.04 

12.24 

14.85 

15.79 

13.46 

13.20 

15.19 

17.30 

305 

13.91 

12.84 

16.10 

17.65 

14.84 

14.47 

17.24 

19.14 

306 

13.85 

12.74 

15.26 

15.60 

13.98 

14.26 

15.65 

17.94 

307 

15.68 

14.97 

15.50 

18.20 

16.41 

16.32 

17.74 

17.80 

308 

13.95 

13.08 

15.94 

16.89 

15.04 

14.94 

16.62 

18.69 

309 

15.97 

14.98 

17.90 

19.01 

16.24 

16.12 

18.17 

20.47 

310 

15.04 

12.96 

16.55 

18.32 

16.07 

15.77 

17.85 

20.47 

311 

14.37 

13.56 

15.99 

17.60 

15.44 

15.34 

17.64 

19.54 

312 

15.37 

14.20 

17.25 

19.25 

16.57 

16.57 

19.50 

22.33 

313 

16.89 

14.91 

17.75 

19.11 

17.80 

17.63 

18.81 

21.03 

314 

18.36 

16.76 

• 

22.29 

20.29 

19.68 

22.29 

24.80 

315 

13.32 

12.06 

14.06 

15.06 

14.23 

14.12 

14.96 

16.92 

316 

13.41 

11.81 

14.73 

15.17 

14.01 

13.68 

15.23 

17.73 

317 

15.10 

14.35 

17.81 

18.83 

16.39 

16.20 

18.94 

21.29 

318 

13.76 

13.20 

15.37 

15.75 

14.22 

13.97 

16.41 

18.51 

319 

12.92 

12.38 

13.54 

14.70 

13.52 

13.16 

14.38 

16.56 

320 

12.65 

11.31 

13.29 

14.25 

13.21 

13.04 

14.23 

15.62 

321 

12.08 

11.18 

13.30 

13.72 

12.56 

12.54 

12.96 

15.54 

322 

14.65 

13.83 

16.64 

17.08 

15.57 

15.30 

16.99 

18.51 

323 

11.15 

10.55 

12.28 

13.20 

11.42 

11.77 

13.67 

14.88 

324 

12.37 

11.12 

13.29 

14.93 

13.20 

13.12 

14.54 

16.42 

325 

10.44 

9.79 

11.53 

11.69 

10.70 

10.77 

11.96 

13.63 

326 

9.31 

8.85 

10.89 

11.23 

9.76 

9.62 

11.22 

12.85 

327 

9.84 

9.05 

10.87 

11.36 

10.05 

10.01 

10.90 

12.66 

328 

9.58 

9.14 

10.81 

10.65 

9.74 

9.88 

10.43 

11.57 

329 

9.26 

8.48 

10.57 

10.09 

9.39 

9.05 

9.68 

12.16 

330 

8.65 

8.19 

9.42 

9.85 

8.91 

8.93 

9.79 

10.82 

331 

9.25 

8.40 

9.61 

10.09 

9.42 

9.38 

9.88 

• 

332 

8.64 

8.03 

9.73 

9.53 

8.70 

8.57 

8.87 

10.64 

333 

8.46 

7.97 

9.26 

9.24 

8.65 

8.68 

9.10 

10.09 

334 

41.96 

37.61 

44.97 

52.64 

46.67 

48.28 

51.65 

54.18 

335 

29.81 

24.80 

29.91 

34.54 

32.13 

30.92 

34.41 

36.89 

336 

23.35 

21.27 

25.65 

29.63 

25.69 

25.71 

28.86 

31.05 


404 


# 

ALIM 

ALIV 

337 

22.66 

19.93 

338 

20.39 

18.76 

339 

18.31 

17.84 

340 

20.43 

17.75 

341 

19.05 

16.94 

342 

18.97 

19.43 

343 

17.76 

16.23 

344 

16.89 

15.10 

345 

16.32 

14.64 

346 

15.61 

14.08 

347 

13.34 

12.08 

348 

11.15 

10.39 

349 

10.87 

10.19 

350 

10.87 

10.43 

351 

10.02 

9.47 

352 

8.87 

8.33 

353 

6.71 

6.17 

354 

30.01 

30.56 

355 

39.61 

39.81 

356 

• 

• 

357 

• 

« 

358 

• 

• 

359 

• 

• 

360 

• 

• 

361 

• 

• 

362 

31.81 

31.01 

363 

• 

• 

364 

29.12 

28.76 

365 

• 

• 

366 

12.15 

11.08 

367 

10.41 

10.22 

368 

10.32 

9.84 

369 

10.21 

9.84 

370 

• 

• 

371 

42.68 

• 

372 

66.42 

67.34 


ALV 

IL 1 

IL 2 

24.56 

27.01 

25.04 

23.37 

24.92 

22.32 

23.17 

24.38 

20.31 

21.60 

22.89 

21.30 

21.28 

23.20 

20.32 

19.75 

23.12 

20.68 

19.11 

21.75 

19.49 

19.03 

19.62 

17.81 

17.24 

18.94 

17.52 

16.69 

18.07 

17.12 

14.47 

15.94 

14.88 

12.35 

12.43 

11.31 

12.01 

11.93 

10.82 

12.22 

11.91 

11.01 

10.99 

11.29 

10.39 

9.62 

9.15 

8.96 

7.52 

7.08 

6.94 

34.02 

31.86 

• 

45.65 

• 

43.20 

• 

• 

39.17 

• 

• 

• 


32.57 

• 

• 

• 

31.80 

37.18 

36.04 

31.43 

14.33 

14.82 

12.42 

12.89 

13.16 

10.52 

12.73 

11.64 

10.26 

12.00 

• 

11.87 

• 

10.83 

• 

• 

79.18 

• 

69.93 

• 

65.16 


IL 3 

IL 4 

IL 5 

23.89 

25.70 

30.05 

22.27 

24.87 

28.01 

19.82 

24.19 

27.50 

20.73 

23.47 

24.67 

20.26 

22.65 

25.16 

20.24 

22.99 

23.99 

19.35 

21.32 

23.68 

17.68 

19.28 

22.45 

17.27 

18.78 

20.91 

16.87 

18.08 

20.07 

14.45 

15.92 

17.94 

11.40 

12.70 

14.39 

11.19 

11.81 

13.51 

10.86 

12.01 

12.57 

10.31 

11.17 

12.18 

8.94 

9.42 

10.54 

6.87 

7.01 

8.49 

29.28 

• 

32.68 

37.93 

• 

• 

• 

• 

• 

• 

• 

44.98 

• 

• 

• 

• 

• 

• 

31.26 

• 

• 

• 

32.75 

• 

• 

• 

33.13 

31.28 

37.57 

45.50 

12.38 

13.85 

17.00 

10.49 

12.01 

16.01 

10.46 

11.69 

14.36 

10.68 

• 

11.92 

• 

13.98 

• 

• 

65.43 

• 

69.43 

• 

72.13 


405 


# 

ALLI 

ALLII 

ALLIII 

ALLIV 

ALLV 

ALWI 

ALWII 

ALWI II 

ALWIV 

1 

10.16 

10.08 

7.06 

9.57 

10.07 

3.20 

3.04 

2.84 

3.00 

2 

5.27 

5.23 

3.72 

5.66 

5.21 

2.40 

2.23 

1.85 

2.38 

3 

3.81 

3.07 

2.03 

2.97 

3.53 

1.89 

1.52 

1.21 

1.58 

4 

2.89 

2.67 

1.71 

2.78 

2.92 

1.47 

1.38 

1.16 

1.35 

5 

2.45 

1.68 

0.97 

1.76 

2.45 

1.14 

0.99 

0.69 

0.97 

6 

• 

• 

• 

• 

• 

0.68 

0.58 

• 

0.57 

7 

8.78 

8.69 

6.59 

8.57 

8.97 

2.80 

2.47 

2.40 

2.79 

8 

5.18 

5.27 

3.46 

4.66 

5.05 

1.62 

1.81 

1.35 

1.63 

9 

3.13 

3.44 

2.61 

3.51 

3.44 

1.61 

1.65 

1.40 

1.68 

10 

3.81 

3.58 

2.01 

3.00 

3.54 

1.79 

1.63 

1.81 

1.57 

11 

3.96 

3.64 

2.29 

3.67 

3.93 

2.24 

1.86 

1.34 

1.94 

12 

3.35 

3.50 

1.61 

3.49 

3.91 

1.39 

1.60 

1.18 

1.63 

13 

2.09 

1.54 

• 

1.53 

2.38 

1.02 

1.65 

• 

0.92 

14 

• 

1.99 

1.91 

2.04 

• 

1.25 

1.07 

0.67 

1.06 

15 

0.88 

1.84 

• 

1.80 

1.37 

0.60 

0.77 

• 

0.73 

16 

2.22 

2.20 

• 

0.87 

• 

1.00 

1.10 

• 

0.54 

17 

9.38 

9.42 

7.45 

9.60 

9.69 

4.03 

3.78 

3.34 

3.70 

18 

5.76 

6.45 

5.48 

6.48 

5.91 

2.20 

2.18 

1.98 

2.00 

19 

8.73 

6.79 

5.29 

6.95 

8.81 

1.98 

2.03 

1.85 

1.73 

20 

8.19 

8.80 

6.95 

8.72 

8.55 

3.53 

3.63 

• 

3.49 

21 

2.65 

2.54 

2.56 

2.65 

2.61 

0.88 

0.97 

0.95 

1.11 

22 

2.41 

2.09 

1.04 

1.87 

2.36 

1.06 

1.11 

0.88 

1.11 

23 

3.77 

3.70 

2.47 

3.42 

3.94 

1.63 

1.65 

1.14 

1.53 

24 

3.59 

2.74 

1.63 

2.95 

3.64 

1.54 

1.43 

1.06 

1.35 

25 

2.87 

. 

2.20 

2.92 

3.07 

1.52 

• 

1.01 

1.52 

26 

• 

3.00 

1.90 

3.20 

• 

1.67 

1.38 

0.96 

1.29 

27 

• 

2.90 

• 

2.87 

• 

• 

1.19 

• 

1.27 

28 

6.85 

7.16 

4.82 

6.98 

6.80 

2.61 

2.59 

1.93 

2.52 

29 

4.83 

4.34 

2.79 

4.43 

4.90 

1.86 

1.73 

1.19 

1.72 

30 

• 

2.46 

1.82 

2.47 

• 

1.29 

1.14 

0.66 

1.07 

31 

. 

2.19 

• 

2.09 

• 

• 

0.96 

• 

1.04 

32 

• 

1.48 

0.96 

1.58 

• 

1.40 

0.92 

0.71 

0.99 

33 

• 

0.78 

• 

0.69 

• 

0.58 

0.39 

• 

0.36 

34 

0.68 

• 

• 

• 

0.62 

0.39 

• 

• 

• 

35 

7.32 

7.13 

5.00 

6.85 

7.37 

3.04 

2.71 

2.28 

2.65 

36 

8.44 

7.27 

5.96 

7.91 

8.31 

2.59 

2.43 

2.13 

2.33 

37 

8.38 

8.03 

5.98 

7.54 

8.28 

2.94 

3.00 

2.19 

3.02 

38 

4.40 

3.42 

2.74 

4.14 

4.34 

1.54 

1.86 

1.14 

1.67 

39 

2.92 

2.46 

1.28 

2.26 

2.84 

1.48 

1.29 

0.86 

1.30 

40 

2.28 

2.19 

• 

2.12 

2.31 

1.16 

1.09 

0.74 

1.16 

41 

2.74 

2.78 

2.26 

2.64 

2.59 

1.21 

1.06 

0.80 

1.19 

42 

2.87 

2.33 

1.37 

2.55 

2.97 

1.48 

1.32 

0.88 

1.42 

43 

4.68 

3.12 

1.72 

3.30 

• 

1.71 

1.52 

1.32 

1.57 

44 

• 

1.54 

1.02 

1.68 

1.73 

• 

0.88 

0.71 

0.96 

45 

• 

1.18 

• 

0.97 

• 

0.90 

0.83 

• 

0.68 

46 

10.88 

9.91 

7.70 

9.41 

9.72 

3.30 

3.30 

2.54 

3.04 

47 

• 

2.38 

• 

2.54 

• 

1.28 

1.09 

• 

1.09 

48 

2.56 

2.19 

1.76 

2.24 

2.89 

1.06 

1.11 

0.71 

1.10 


406 


# 

ALU 

ALLII 

ALUM 

ALLIV 

49 

• 

1.93 

• 

1.01 

50 

• 

1.80 

• 

1.34 

51 

• 

1.35 

1.21 

• 

52 

• 

1.23 

• 

1.15 

53 

• 

• 

• 

• 

54 

• 

• 

• 

• 

55 

• 

1.42 

• 

1.32 

56 

• 

0.66 

• 

• 

57 

• 

• 

• 

• 

58 

• 

4.41 

2.97 

4.55 

59 

4.59 

3.88 

2.90 

3.97 

60 

7.87 

6.73 

4.77 

7.40 

61 

3.81 

3.46 

2.28 

3.50 

62 

2.26 

1.91 

1.84 

1.95 

63 

3.60 

4.19 

3.02 

3.73 

64 

3.67 

3.61 

1.75 

3.36 

65 

2.89 

2.47 

2.46 

2.69 

66 

9.63 

9.43 

7.18 

9.06 

67 

3.41 

3.87 

2.74 

3.83 

68 

4.31 

3.98 

2.00 

3.68 

69 

9.34 

7.74 

4.41 

7.41 

70 

2.97 

3.45 


3.31 

71 

4.87 

4.41 

2.75 

4.34 

72 


2.37 

1.48 

2.69 

73 

7.16 

8.53 

5.88 

8.31 

74 

3.31 

2.67 

1.77 

2.69 

75 

2.41 

2.09 

1.51 

2.13 

76 

3.81 

3.91 

2.97 

3.46 

77 

10.00 

10.49 

7.44 

8.59 

78 


2.13 

1.62 

1.98 

79 

• 

2.37 

1.24 

2.29 

80 

• 

1.77 

1.85 

2.00 

81 

0.66 

1.47 

• 

1.38 

82 

2.05 

1.98 

• 

1.96 

83 

1.72 

1.42 

0.52 

1.65 

84 

1.53 

1.27 

• 

1.01 

85 

• 

• 

2.65 

• 

86 

2.71 

2.71 

1.73 

2.54 

87 

• 

3.07 

1.42 

2.99 

88 

2.00 

2.71 

2.81 

2.78 

89 

7.95 

9.56 

7.36 

9.24 

90 

• 

1.79 

1.10 

• 

91 

2.76 

2.27 

1.32 

2.27 

92 

• 

2.13 

1.84 

1.84 

93 

• 

1.99 

• 

2.43 

94 

2.76 

1.86 

1.58 

1.98 

95 

• 

1.43 

• 

1.43 

96 

2.13 

1.73 

1.27 

1.79 


ALLV 

ALWI 

ALWII 

ALWI II 

ALWIV 

• 

0.67 

1.11 

• 

0.58 

• 

0.67 

0.57 

• 

0.88 

• 

• 

0.69 

0.54 

• 

1.07 

0.86 

0.58 

• 

0.66 

• 

0.48 

0.46 

• 

0.50 

• 

0.41 

0.48 

• 

0.53 

• 

• 

0.53 

• 

0.63 

5.41 

. 

1.96 

1.33 

1.79 

4.31 

1.70 

1.42 

1.46 

1.42 

• 

2.29 

2.24 

1.67 

2.23 

3.97 

1.90 

1.75 

1.33 

1.75 

1.91 

1.11 

0.90 

0.83 

1.04 

3.93 

1.52 

1.53 

1.23 

1.61 

3.82 

1.82 

1.63 

1.09 

1.72 

2.99 

1.52 

1.33 

1.29 

1.43 

10.66 

4.15 

3.98 

2.89 

3.81 

3.72 

1.33 

1.46 

1.25 

1.34 

4.21 

2.04 

1.94 

1.21 

1.87 

8.72 

2.87 

2.89 

2.19 

3.12 

« 

1.43 

1.82 

1.23 

1.87 

4.72 

2.20 

1.77 

1.27 

1.82 

• 

1.23 

1.24 

0.88 

1.24 

• 

3.12 

2.98 

2.31 

2.89 

3.22 

1.44 

1.24 

0.80 

1.14 

2.28 

1.06 

1.01 

0.77 

1.09 

3.37 

2.10 

2.08 

1.80 

2.15 

10.24 

3.30 

3.68 

3.23 

3.88 

• 

• 

0.95 

0.74 

0,97 

« 

1.94 

1.62 

0.96 

1.58 

• 

1.14 

0.96 

0.77 

1.09 

1.00 

0.63 

0.63 

• 

0.66 

2.28 

1.18 

1.10 

• 

1.07 

• 

0.74 

0.81 

0.43 

0.86 

1.82 

0.93 

0.72 

• 

0.72 

• 

• 

• 

1.06 

1.10 

• 

1.54 

1.39 

1.20 

1.48 

• 

• 

1.54 

0.91 

1.51 

2.45 

1.10 

1.34 

1.37 

1.29 

8.22 

3.55 

3.27 

2.92 

2.97 

• 

1.28 

0.91 

0.57 

0.76 

2.62 

1.10 

1.04 

0.77 

0.97 

• 

• 

1.06 

0.78 

1.01 

• 

0.82 

0.81 

• 

0,86 

2.69 

1.14 

1.09 

0.66 

1.09 

• 

1.25 

1.04 

• 

1.07 

2.18 

0.87 

0.83 

0.49 

0.82 


407 


# 

ALLI 

ALLII 

ALLIII 

ALLIV 

97 

0.88 

1.51 

• 

1.52 

98 

9.04 

8.94 

7.97 

8.87 

99 

6.19 

4.55 

4.76 

• 

100 

8.52 

9.01 

6.60 

* 

101 

6.66 

6.56 

5.18 

6.60 

102 

7.48 

7.08 

4.68 

6.89 

103 

• 

2.81 

1.76 

2.92 

104 

1.52 

1.68 

• 

1.81 

105 

• 

1.23 

0.86 

1.65 

106 

• 

1.68 

• 

1.77 

107 

2.48 

2.29 

1.62 

2.14 

108 

2.03 

1.62 

0.86 

1.30 

109 

1.54 

1.29 

0.88 

1.09 

110 

3.78 

3.03 

2.14 

3.02 

111 

• 

1.29 

• 

1.28 

112 

1.49 

1.35 

1.01 

1.49 

113 

0.95 

1.11 

• 

• 

114 

• 

1.38 

0.63 

1.37 

115 

1.14 

1.32 

• 

1.32 

116 

2.52 

1.93 

1.15 

1.80 

117 

• 

• 

• 

• 

118 

• 

• 

• 

• 

119 

• 

• 

• 

• 

120 

5.42 

7.40 

4.74 

7.25 

121 

4.44 

4.29 

2.85 

4.01 

122 

2.08 

2.18 

• 

2.08 

123 

1.53 

1.44 

1.02 

1.48 

124 

0.71 

0.66 

• 

0.68 

125 

1.13 

1.21 

0.86 

1.00 

126 

• 

• 

• 

• 

127 

10.16 

10.05 

7.87 

9.93 

128 

2.36 

2.24 

2.01 

2.36 

129 

8.66 

8.83 

7.06 

8.63 

130 

17.83 

17.28 

12.59 

15.97 

131 

5.85 

• 

3.04 

4.85 

132 

7.46 

7.01 

6.19 

6.90 

133 

5.84 

5.63 

5.18 

• 

134 

• 

6.95 

4.54 

7.21 

135 

6.47 

6.13 

4.88 

6.21 

136 

4.50 

4.74 

3.55 

4.22 

137 

0.92 

1.13 

• 

0.87 

138 

8.45 

7.84 

6.18 

7.42 

139 

6.47 

6.21 

4.99 

5.84 

140 

6.21 

5.20 

3.49 

5.47 

141 

0.99 

0.40 

• 

1.13 

142 

• 

5.71 

3.60 

5.39 

143 

• 

• 

• 

• 

144 

1.34 

1.75 

• 

1.87 


ALLV 

ALWI 

ALWII 

ALWI II 

ALWIV 

1.54 

0.58 

0.62 

• 

0.73 

8.82 

3.32 

3.02 

2.73 

2.79 

6.08 

2.47 

2.24 

2.03 

2.51 

8.85 

2.84 

2.64 

2.27 

2.61 

6.68 

1.87 

1.76 

1.75 

1.93 

7.62 

2.09 

2.31 

2.03 

2.42 

. 

1.80 

1.49 

0.97 

1.37 

• 

0.86 

0.82 

• 

0.86 

• 

1.28 

0.76 

0.34 

0.86 

• 

0.96 

0.77 

• 

0.77 

2.59 

1.20 

1.01 

0.81 

0.97 

2.10 

1.06 

0.95 

0.58 

0.87 

1.23 

0.52 

0.55 

0.39 

0.55 

3.84 

1.52 

1.33 

0.95 

1.29 

« 

0.97 

0.82 

• 

0.78 

1.94 

0.80 

0.73 

0.63 

0.78 

• 

0.74 

0.72 

• 

0.59 

1.58 

• 

0.91 

0.63 

0.95 

1.21 

0.71 

0.73 

• 

0.91 

2.61 

1.20 

1.09 

0.76 

1.16 

• 

• 

0.41 

• 

0.45 

• 

• 

• 

0.57 

• 

6.89 

• 

2.41 

• 

2.29 

• 

2.20 

• 

2.23 

4.38 

1.91 

1.94 

1.52 

1.95 

2.22 

1.20 

1.14 

• 

1.21 

1.42 

0.62 

0.66 

0.43 

0.77 

0.64 

0.52 

0.52 

0.45 

0.58 

1.09 

0.88 

0.66 

0.46 

0.81 

10.50 

3.92 

4.14 

3.78 

3.88 

2.09 

1.29 

1.37 

0.96 

1.43 

9.22 

2.87 

2.75 

2.99 

2.92 

16.51 

5.30 

5.49 

4.22 

5.60 

6.05 

2.43 

2.13 

1.93 

2.18 

7.55 

1.93 

1.87 

1.81 

1.98 

6.35 

2.10 

2.36 

2.23 

• 

. 

• 

1.95 

1.85 

2.14 

• 

2.46 

2.50 

2.12 

2.34 

4.83 

2.08 

1.98 

1.94 

2.23 

0.91 

0.46 

0.53 

• 

0.48 

8.76 

2.00 

1.87 

1.72 

1.85 

6.45 

1.73 

1.73 

1.53 

1.95 

6.14 

2.36 

2.43 

1.66 

2.13 

1.09 

0.59 

0.25 

• 

0.66 

7.78 

• 

2.09 

1.60 

1.99 

• 

2.87 

3.51 

3.41 

2.33 

0.99 

0.59 

0.99 

• 

1.06 


408 


# 

ALU 

145 

1.49 

146 

• 

147 

6.87 

148 

7.07 

149 

12.94 

150 

11.92 

151 

12.28 

152 

17.15 

153 

9.20 

154 

14.92 

155 

10.12 

156 

6.56 

157 

7.39 

158 

9.65 

159 

10.71 

160 

14.13 

161 

8.11 

162 

11.17 

163 

11.76 

164 

9.51 

165 

14.14 

166 

15.80 

167 

9.05 

168 

12.44 

169 

9.74 

170 

8.97 

171 

5.15 

172 

15.46 

173 

13.14 

174 

9.25 

175 

10.80 

176 

9.18 

177 

• 

178 

11.58 

179 

11.51 

180 

11.36 

181 

9.97 

182 

8.24 

183 

11.38 

184 

9.81 

185 

10.42 

186 

9.66 

187 

10.66 

188 

11.96 

189 

13.69 

190 

13.39 

191 

10.60 

192 

10.99 


ALLII ALLIII 

1.80 
1.68 

7.03 

6.99 5.44 

12.83 
10.97 
12.42 

15.96 
9.35 

12.36 

12.69 

6.49 
7.79 
10.88 

11.03 

13.54 

8.92 

9.54 
10.85 

9.96 

12.92 
14.59 
9.38 
11.40 
7.81 

8.46 

15.26 

11.30 

9.34 

9.09 

11.36 

8.54 

• • 

9.55 
9.71 

8.99 

8.69 

10.44 
9.84 
9.06 

9.49 

9.83 

10.45 
11.97 

13.36 
11.72 
9.19 


ALLIV 

ALLV 

2.03 

1.73 

8.14 

8.30 

• 

7.20 

11.77 

9.51 

10.87 

11.49 

13.10 

13.15 

16.50 

15.55 

8.71 

8.43 

13.09 

12.92 

11.80 

11.30 

6.21 

6.93 

• 

7.68 

10.57 

10.65 

10.84 

10.49 

12.75 

13.47 

9.38 

10.35 

10.80 

11.48 

10.81 

12.08 

9.86 

10.41 

12.65 

13.31 

13.36 

15.75 

9.84 

10.38 

11.21 

13.43 

9.10 

9.49 

9.43 

9.11 

8.08 

7.78 

13.71 

14.67 

11.43 

11.59 

8.83 

9.84 

9.25 

10.12 

10.59 

10.23 

8.48 

9.79 

11.83 

11.51 

9.69 

12.10 

9.85 

10.75 

• 

9.95 

8.84 

8.19 

10.26 

11.32 

11.04 

10.56 

8.30 

10.76 

9.44 

9.82 

10.32 

11.35 

10.35 

12.83 

13.03 

12.11 

13.52 

13.67 

11.26 

12.00 

9.89 

11.44 


ALWI 

ALWII 

0.91 

1.01 

1.47 

0.95 

2.69 

2.92 

1.68 

1.76 

2.55 

2.38 

2.47 

2.45 

1.94 

1.87 

3.01 

2.91 

2.08 

2.04 

2.28 

2.55 

2.51 

2.41 

1.42 

1.62 

2.17 

2.26 

2.73 

2.60 

1.98 

2.14 

2.88 

2.72 

2.31 

2.46 

1.99 

1.94 

2.37 

2.61 

2.04 

2.12 

2.78 

2.84 

2.75 

3.06 

1.98 

2.07 

2.47 

2.46 

1.73 

2.01 

2.21 

• 

2.07 

2.31 

2.07 

2.43 

2.70 

2.73 

1.83 

1.95 

1.87 

1.94 

2.36 

2.42 

• 

2.23 

2.57 

• 

2.11 

2.06 

2.32 

2.22 

2.11 

2.35 

1.75 

2.00 

1.86 

2.35 

1.89 

1.97 

2.04 

2.25 

1.64 

2.11 

2.49 

2.56 

2.56 

2.25 

2.43 

2.37 

2.50 

2.27 

2.24 

2.37 

2.18 

2.27 


ALWIII ALWIV 

1.09 

• • 

2.76 2.59 

1.70 

2.41 

2.52 

1.90 

• 2.96 

2.09 

• 2.55 
2.79 
1.71 
2.27 
2.78 
2.51 

2.91 

2.32 
2.01 

2.33 

2.17 
2.86 

2.89 
2.50 

2.54 

2.03 
2.21 
2.30 
2.37 
2.75 

2.04 

1.90 

• 2.33 

• 2.51 
2.93 

• 2.14 

• 2.14 

• 2.11 
1.90 
2.33 

2.03 

2.18 
1.96 
2.64 

2.45 

2.55 
2.36 
2.24 

• 2.32 


409 


# 

ALU 

ALLIl 

ALLIII 

ALLIV 

193 

10.19 

9.23 

« 

9.28 

194 

6.01 

5.68 

• 

5.28 

195 

8.23 

8.22 

• 

7.89 

196 

9.35 

10.05 

• 

9.67 

197 

12.31 

11.68 

• 

12.76 

198 

8.05 

8.85 

• 

9.32 

199 

11.78 

9.72 

• 

10.38 

200 

7.97 

7.13 

• 

7.69 

201 

8.75 

8.73 

• 

8.12 

202 

10.29 

11.23 

• 

10.47 

203 

8.99 

9.22 

• 

8.46 

204 

16.35 

16.17 

• 

14.42 

205 

6.26 

5.86 

• 

5.62 

206 

15.79 

14.04 

• 

15.04 

207 

9.56 

9.25 

• 

9.56 

208 

• 

9.65 

• 

10.02 

209 

6.52 

• 

• 

5.52 

210 

7.21 

7.60 

• 

5.75 

211 

10.55 

9.62 

• 

9.65 

212 

10.33 

10.37 

• 

9.73 

213 

8.14 

9.26 

• 

9.02 

214 

12.38 

12.13 

• 

11.94 

215 

8.31 

7.70 

• 

7.05 

216 

5.88 

6.42 


8.51 

217 

8.73 

9.35 


8.58 

218 

11.82 

11.40 


12.12 

219 

12.26 

11.33 


11.74 

220 

9.14 

9.07 

• 

9.05 

221 

10.04 

10.05 


10.15 

222 

11.93 

12.09 

• 

11.29 

223 

16.64 

13.76 

• 

15.08 

224 

• 

19.88 


17.04 

225 

• 

14.05 

• 

14.07 

226 

8.66 

8.72 


8.67 

227 

28.10 


• 

26.70 

228 

28.23 

27.71 

• 

27.12 

229 

20.05 

17.36 

• 

16.34 

230 

15.25 

12.05 

t 

12.73 

231 

23.38 

20.39 

• 

20.24 

232 

17.60 

15.77 

« 

15.58 

233 

10.17 

6.19 

3.21 

6.01 

234 

10.82 

7.22 

3.81 

6.09 

235 

6.95 

4.50 

3.18 

4.36 

236 

9.81 

4.58 

2.98 

5.05 

237 

7.95 

4.27 

3.28 

4.90 

238 

8.62 

4.30 

2.51 

4.45 

239 

6.38 

2.79 

2.27 

2.60 

240 

7.55 

5.81 

3.61 

5.32 


ALLV 

ALWI 

ALWII 

ALWI II 

ALWIV 

10.68 

2.65 

2.55 

• 

2.57 

5.52 

1.30 

1.80 

• 

1.71 

8.47 

2.28 

2.32 

• 

2.27 

9.62 

2.71 

• 

• 

2.67 

12.30 

2.31 

2.50 

• 

2.62 

8.72 

2.19 

2.42 

• 

2.78 

11.83 

2.48 

2.32 

• 

2.36 

8.00 

1.91 

2.38 

• 

2.01 

9.48 

1.98 

1.70 

• 

1.92 

9.60 

2.57 

2.51 

• 

2.41 

10.05 

1.95 

1.91 

• 

2.09 

16.95 

2.88 

2.80 

• 

2.89 

8.24 

1.61 

1.70 

• 

1.68 

16.40 

2.96 

2.71 

• 

2.85 

9.66 

2.26 

2.05 

• 

2.15 

12.48 

• 

2.36 

• 

2.27 

5.95 

1.69 

• 

• 

1.72 

9.38 

1.36 

1.71 

• 

1.75 

11.84 

2.29 

2.22 

• 

2.25 

10.44 

2.12 

2.04 

• 

2.34 

8.30 

2.04 

2.19 

• 

2.09 

11.31 

2.38 

2.18 

• 

2.29 

8.68 

1.91 

1.76 

• 

1.99 

8.55 

1.86 

1.93 

• 

2.06 

8.82 

2.10 

2.17 

• 

2.24 

12.84 

2.80 

2.57 

• 

2.64 

10.16 

1.98 

1.83 

• 

1.82 

8.56 

1.41 

1.82 

• 

1.72 

9.92 

2.21 

2.21 

• 

2.08 

12.71 

2.52 

2.52 

• 

2.51 

17.52 

2.09 

2.50 

2.26 

2.28 

23.07 

• 

3.40 

• 

• 

17.53 

• 

2.47 

• 

2.52 

9.57 

2.52 

2.57 

• 

2.48 

29.46 

3.23 

• 

• 

3.34 

27.90 

2.93 

2.73 

• 

2.59 

20.12 

3.00 

3.14 

3.06 

3.12 

16.85 

2.29 

2.60 

• 

2.59 

23.48 

2.31 

3.25 

• 

3.08 

17.53 

2.05 

2.14 

• 

2.04 

11.23 

3.84 

3.39 

1.61 

3.28 

10.65 

4.71 

3.59 

• 

3.49 

7.97 

4.07 

2.90 

• 

2.80 

9.57 

5.71 

3.77 

• 

3.06 

8.86 

4.40 

2.75 

• 

3.08 

8.73 

4.20 

2.88 

• 

3.18 

6.42 

3.28 

2.60 

• 

2.20 

8.45 

4.25 

3.75 

• 

3.47 


410 


# 

ALU 

ALLII 

ALLIII 

ALLIV 

241 

6.62 

4.24 

2.42 

4.02 

242 

5.30 

3.12 

1.29 

3.20 

243 

4.92 

3.86 

1.96 

2.97 

244 

5.21 

3.25 

1.27 

2.59 

245 

4.17 

2.05 

1.27 

2.15 

246 

4.85 

2.69 

1.39 

3.16 

247 

4.62 

2.51 

1.28 

2.43 

248 

4.01 

2.09 

1.10 

1.89 

249 

4.80 

3.26 

1.06 

3.44 

250 

4.59 

2.29 

0.92 

2.37 

251 

4.49 

2.29 

0.71 

2.24 

252 

• 

• 

0.64 

1.85 

253 

4.47 

2.54 

0.86 

2.10 

254 

2.48 

0.78 

• 

1.04 

255 

3.55 

1.49 

0.88 

1.63 

256 

4.14 

1.43 

• 

1.76 

257 

3.04 

1.63 

0.63 

1.73 

258 

3.41 

1.91 

• 

2.26 

259 

3.18 

1.48 

0.90 

1.15 

260 

3.06 

1.37 

• 

1.58 

261 

3.14 

1.54 

0.45 

1.51 

262 

2.18 

1.15 

• 

1.29 

263 

7.51 

4.50 

3.92 

4.06 

264 

6.63 

2.68 

3.57 

5.28 

265 

8.10 

3.34 

2.88 

3.58 

266 

11.21 

5.57 

3.01 

6.89 

267 

13.21 

9.24 

6.60 

10.64 

268 

12.75 

9.26 

6.62 

8.64 

269 

9.76 

7.69 

6.64 

7.25 

270 

14.54 

7.18 

5.10 

6.11 

271 

12.24 

7.39 

5.61 

6.62 

272 

8.66 

5.26 

5.13 

5.01 

273 

7.63 

4.88 

2.34 

4.11 

274 

7.11 

5.42 

3.91 

4.38 

275 

8.06 

5.40 

4.66 

4.86 

276 

5.76 

3.88 

2.38 

3.85 

277 

8.95 

5.44 

4.06 

5.86 

278 

5.92 

3.90 

3.32 

4.74 

279 

4.78 

4.28 

• 

4.05 

280 

8.95 

5.10 

3.44 

5.60 

281 

6.18 

3.74 

4.02 

4.64 

282 

9.66 

5.66 

4.14 

6.11 

283 

8.55 

4.91 

3.84 

5.13 

284 

6.71 

3.91 

3.26 

3.87 

285 

4.63 

2.86 

2.33 

3.06 

286 

8.14 

4.39 

3.13 

4.03 

287 

6.86 

4.11 

3.53 

3.61 

288 

5.70 

3.54 

3.14 

3.22 


ALLV 

ALWI 

ALWII 

ALWI II 

ALWIV 

6.43 

3.41 

2.80 

• 

2.79 

5.58 

3.44 

2.38 

• 

2.57 

5.80 

2.78 

2.94 

• 

2.22 

5.16 

3.20 

2.28 

• 

2.20 

4.55 

2.92 

1.48 

• 

1.58 

5.18 

3.11 

1.79 

• 

2.10 

4.54 

2.52 

1.98 

• 

1.86 

5.20 

2.40 

1.77 

• 

1.73 

5.53 

2.66 

1.76 

• 

2.00 

4.66 

2.65 

1.80 

• 

1.98 

4.26 

2.59 

2.15 

• 

2.28 

• 

• 

• 

• 

1.48 

4.20 

2.62 

2.18 

• 

1.76 

3.47 

1.73 

• 

• 

• 

4.49 

2.08 

1.81 


1.21 

4.01 

2.36 

1.46 

• 

1.67 

3.25 

1.94 

1.58 

« 

1.71 

3.31 

2.28 

1.51 


1.73 

3.16 

2.19 

1.40 

• 

1.40 

3.47 

2.08 

1.43 

• 

1.54 

2.57 

2.05 

1.63 


1.47 

2.57 

1.43 


• 

• 

6.66 

4.30 

3.39 

2.81 

3.57 

7.07 

3.64 

2.99 

2.48 

2.74 

7.39 

3.37 

2.14 

• 

2.15 

12.45 

4.67 

4.16 

2.97 

3.98 

14.54 

7.37 

5.02 

3.59 

4.48 

13.03 

7.30 

5.24 

5.62 

5.61 

13.03 

6.36 

4.67 

4.36 

5.05 

13.06 

5.48 

3.47 

3.24 

3.57 

12.53 

5.80 

4.51 

3.59 

3.97 

9.15 

3.91 

3.05 

2.52 

3.36 

8.91 

4.12 

3.24 

2.67 

3.45 

7.92 

3.13 

3.06 

2.92 

3.31 

8.52 

4.88 

3.28 

3.12 

3.80 

5.53 

2.65 

2.21 

• 

2.32 

• 

4.10 

3.42 

2.80 

3.05 

5.85 

3.01 

2.95 

• 

2.93 

5.93 

3.06 

2.66 

• 

2.24 

9.07 

3.95 

2.57 

2.42 

2.82 

6.71 

3.14 

2.37 

• 

2.61 

10.06 

5.00 

3.27 

3.21 

3.67 

9.34 

3.77 

2.67 

• 

2.65 

6.42 

2.89 

2.52 

• 

2.40 

4.17 

2.95 

2.11 

• 

2.27 

7.81 

3.77 

2.42 

2.74 

2.65 

7.13 

3.08 

2.52 

• 

2.79 

5.04 

2.83 

2.63 

2.88 

2.66 


411 


# 

ALLI 

ALLII 

ALLIII 

ALLIV 

289 

5.88 

3.35 

3.03 

3.49 

290 

6.62 

4.47 

3.27 

4.03 

291 

5.62 

3.97 

3.56 

3.86 

292 

4.55 

3.64 

3.06 

2.95 

293 

5.87 

3.52 

• 

2.97 

294 

6.28 

3.73 

2.70 

3.68 

295 

6.65 

3.68 

3.11 

4.12 

296 

5.21 

3.98 

2.88 

2.55 

297 

4.16 

3.18 

• 

2.71 

298 

5.56 

3.46 

2.83 

2.98 

299 

4.82 

3.72 

2.24 

3.01 

300 

4.15 

3.13 

2.97 

3.01 

301 

5.07 

2.53 

2.18 

2.75 

302 

5.01 

2.77 

2.08 

2.58 

303 

5.56 

3.47 

2.48 

3.02 

304 

4.71 

2.13 

• 

2.03 

305 

5.00 

2.90 

2.22 

2.64 

306 

4.32 

2.57 

• 

2.44 

307 

3.88 

2.10 

• 

2.04 

308 

4.16 

2.62 

2.38 

3.08 

309 

4.31 

3.22 

• 

2.60 

310 

4.87 

3.09 

2.30 

3.12 

311 

4.44 

2.17 

1.99 

2.50 

312 

5.41 

3.06 

2.76 

3.01 

313 

3.61 

2.79 

• 

2.75 

314 

5.94 

3.53 

3.59 

3.42 

315 

4.44 

2.48 

2.09 

2.32 

316 

4.28 

2.68 

2.16 

2.55 

317 

4.81 

3.36 

2.33 

2.91 

318 

• 

2.97 

2.20 

2.54 

319 

4.05 

2.69 

2.55 

2.61 

320 

3.99 

3.10 

1.89 

2.82 

321 

3.99 

3.04 

• 

2.35 

322 

4.33 

2.41 

2.00 

2.56 

323 

3.89 

2.34 

• 

2.00 

324 

4.11 

2.54 

1.93 

2.09 

325 

3.43 

1.95 

• 

2.15 

326 

3.22 

2.03 

1.53 

1.69 

327 

3.41 

2.07 

1.52 

1.93 

328 

2.89 

1.98 

1.39 

1.83 

329 

2.57 

1.71 

• 

1.24 

330 

2.74 

2.00 

• 

1.64 

331 

2.78 

2.03 

• 

1.86 

332 

2.65 

1.62 

• 

1.77 

333 

2.68 

1.82 

• 

1.52 

334 

12.01 

8.54 

6.58 

8.61 

335 

7.92 

4.92 

3.86 

4.31 

336 

8.21 

4.99 

3.57 

5.05 


ALLV 

ALWI 

ALWII 

ALWI II 

ALWIV 

6.22 

3.28 

2.22 

2.08 

2.43 

6.85 

3.54 

2.87 

• 

2.68 

5.42 

3.21 

2.57 

• 

2.60 

5.56 

2.98 

2.55 

• 

2.80 

5.51 

3.26 

2.41 

• 

2.37 

5.28 

3.24 

2.49 

• 

2.42 

6.74 

3.92 

2.65 

• 

2.87 

5.47 

3.23 

2.41 

• 

2.25 

4.46 

2.04 

2.09 

• 

2.21 

5.95 

3.60 

2.50 


2.46 

5.22 

2.63 

2.73 

• 

2.51 

4.92 

2.59 

2.28 

• 

2.36 

4.79 

2.87 

2.08 

• 

1.97 

4.52 

3.00 

1.97 

• 

1.85 

4.85 

2.58 

2.03 

• 

2.11 

3.30 

2.86 

1.56 

• 

1.49 

5.12 

3.24 

1.93 

• 

2.13 

4.31 

2.34 

1.84 

• 

1.78 

4.10 

2.57 

2.12 

• 

2.28 

4.23 

2.53 

1.71 

1.78 

1.87 

4.64 

2.97 

1.72 

• 

2.10 

4.83 

2.95 

2.05 

• 

2.04 

4.65 

2.40 

1.88 

• 

2.09 

5.32 

2.77 

1.85 

• 

1.98 

4.25 

2.66 

1.85 

• 

2.20 

5.84 

3.48 

2.52 

• 

2.74 

4.24 

2.83 

1.72 

• 

1.72 

3.99 

2.32 

1.63 

• 

1.55 

4.72 

2.59 

1.90 

• 

2.01 

5.16 

2.64 

1.59 

• 

1.55 

4.63 

2.45 

1.98 

• 

1.84 

3.79 

1.94 

1.38 

• 

1.59 

3.44 

2.13 

• 

• 

1.83 

4.06 

2.44 

1.85 

• 

1.80 

3.61 

2.53 

1.50 

• 

1.72 

4.20 

2.44 

1.50 

• 

1.62 

2.83 

2.05 

1.31 

• 

1.59 

2.99 

1.87 

1.07 

• 

1.40 

3.51 

2.19 

1.46 

• 

1.70 

2.80 

1.87 

1.33 

• 

1.35 

2.60 

1.81 

1.20 

• 

1.02 

2.57 

1.94 

• 

• 

1.54 

2.66 

2.00 

1.86 

• 

1.75 

2.49 

1.72 

• 

• 

• 

2.17 

1.85 

• 

• 

• 

12.07 

4.79 

4.76 

4.98 

4.62 

7.09 

3.60 

2.99 

2.89 

3.04 

7.88 

4.75 

3.28 

2.71 

3.45 


412 


# 

ALU 

ALLII 

ALUM 

ALLIV 

337 

7.30 

5.12 

3.72 

4.30 

338 

7.18 

4.17 

3.58 

4.08 

339 

5.79 

4.06 

2.64 

3.40 

340 

6.50 

4.23 

3.29 

4.15 

341 

• 

3.43 

2.88 

3.02 

342 

6.92 

3.62 

3.42 

3.41 

343 

6.71 

4.19 

3.33 

3,97 

344 

5.34 

3.44 

3.17 

3.39 

345 

5,14 

3.60 

2,73 

2.94 

346 

4.13 

2.92 

2.60 

2.82 

347 

4.23 

3.06 

2.58 

2.83 

348 

3,53 

1.98 

• 

2.41 

349 

3.62 

2.27 

• 

2.47 

350 

2.73 

1.67 

• 

1.63 

351 

2.79 

2.04 

• 

2.12 

352 

2.32 

• 

• 

• 

353 

1,81 

• 

• 

• 

354 

5.60 

• 

• 

5.57 

355 

• 

• 

• 

• 

356 

5.48 

5.02 

5.47 

• 

357 

8.05 

7.42 

• 

• 

358 

6.97 

• 

7.15 

• 

359 

• 

5.04 


6.41 

360 

• 

• 

• 

• 

361 

• 

8,29 


• 

362 

5.41 

4.92 

4.25 

4.50 

363 

4.80 

3.39 


• 

364 

• 

• 


• 

365 


• 


• 

366 

• 

• 

• 

• 

367 

• 

• 


• 

368 

• 

• 



369 

• 

• 

• 

• 

370 

• 

• 

• 

• 

371 

13.76 

13.10 

10.40 

• 

372 

15.76 

13.53 

11.20 

12.97 


ALLV 

ALWI 

ALWII 

ALWI II 

ALWIV 

8.04 

3.75 

3.04 

2.77 

2.85 

7.72 

3.34 

2.46 

2.31 

2.41 

6.41 

3.01 

2,31 

2.13 

2.66 

6.62 

3.03 

2.49 

2.59 

2.24 

5.93 

2.60 

1.87 

2.02 

2.17 

6.61 

3.16 

2.77 

2.97 

3.19 

7.16 

3,25 

1.92 

2.27 

2.21 

4.64 

2.52 

2.13 

2.23 

1.92 

5.19 

3.27 

2.06 

1.81 

2.25 

4.29 

2.17 

1.87 

• 

1.97 

3.79 

2.40 

2.33 

2.47 

2.24 

3.65 

2.83 

1.74 

• 

1.85 

3.02 

1.91 

1.54 

• 

1.19 

2.74 

1.96 


• 

• 

2.81 

1.94 

1.98 

• 

1.47 

2.19 

1.50 

• 

• 

• 

1.76 

1.36 

• 


• 

5.14 

2.61 


• 

3.45 

9.57 

• 

• 

• 

• 

• 

2.78 

2.70 

2.52 

• 

• 

4.52 

4.71 


• 

7.55 

4.16 

4.14 

3.97 

• 

4.69 

• 

3.23 

• 

3.56 

9.35 

• 

4.73 

• 

• 

• 

3.37 

3.03 

2.45 

2.80 

4.90 

• 

• 

2.34 

• 

• 

• 

• 

• 

• 

• 

• 

• 

• 

• 

• 

• 

• 

« 

• 

• 

• 

3.08 

• 

• 

• 

3.16 

• 

• 

• 

2.98 

• 

• 

• 

4.25 

15.22 

3.06 

2.59 

2.54 

3.03 


# 

1 

2 

3 

4 

5 

6 

7 

8 

9 

10 

11 

12 

13 

14 

15 

16 

17 

18 

19 

20 

21 

22 

23 

24 

25 

26 

27 

28 

29 

30 

31 

32 

33 

34 

35 

36 

37 

38 

39 

40 

41 

42 

43 

44 

45 

46 

47 

48 


413 


ALWV 

ALPI 

ALPII 

ALPIII 

ALPIV 

ALPV 

3.07 

30.32 

23.93 

23.02 

23.93 

29.99 

2.20 

17.25 

13.88 

15.08 

14.13 

17.57 

1.77 

13.56 

10.92 

11.50 

10.87 

13.58 

1.40 

12.52 

10.73 

11.09 

10.47 

12.54 

1.20 

10.93 

9.76 

9.76 

9.48 

10.68 

0.66 

9.84 

8.73 

• 

8.62 

9.43 

2.67 

28.51 

22.37 

22.42 

22.58 

28.15 

1.58 

17.33 

14.24 

14.03 

14.47 

17.58 

1.63 

17.86 

15.03 

14.83 

14.00 

17.46 

1.66 

18.36 

14.35 

14.43 

14.38 

17.97 

2.18 

15.03 

12.72 

12.94 

12.91 

15.20 

1.56 

14.50 

11.77 

12.39 

12.16 

14.61 

1.19 

12.45 

10.87 

• 

10.88 

12.29 

1.24 

11.77 

9.65 

10.21 

9.76 

11.39 

0.62 

9.81 

8.19 

• 

7.92 

9.08 

• 

15.55 

13.67 

• 

12.86 

15.15 

3.98 

29.93 

23.49 

22.60 

23.76 

29.92 

2.08 

24.85 

21.25 

21.42 

21.22 

25.52 

2.07 

24.05 

18.92 

20.44 

20.38 

23.58 

3.70 

29.51 

24.38 

24.28 

23.85 

29.00 

0.90 

14.98 

12.91 

13.08 

12.68 

14.85 

1.15 

12.09 

9.33 

9.60 

9.16 

11.73 

1.62 

13.19 

10.52 

11.54 

10.71 

13.30 

1.56 

14.32 

11.54 

12.01 

11.48 

14.07 

1.52 

16.20 

• 

14.51 

14.03 

16.61 

• 

12.86 

11.07 

11.83 

10.90 

• 

1.27 

• 

11.86 

• 

11.76 

13.46 

2.54 

21.92 

17.55 

18.42 

17.71 

21.51 

1.75 

15.59 

12.90 

13.86 

13.04 

15.69 

• 

14.18 

12.26 

12.47 

12.01 

• 

1.21 

12.42 

11.27 

• 

10.43 

12.31 

1.40 

10.13 

8.59 

8.76 

8.39 

9.94 

0.57 

9.01 

8.07 

• 

8.14 

8.95 

0.36 

8.71 

• 

• 

• 

8.83 

2.88 

25.80 

20.37 

20.37 

19.74 

26.28 

2.43 

21.79 

18.75 

19.13 

18.59 

22.25 

2.89 

24.56 

20.56 

21.25 

20.80 

24.49 

1.63 

12.50 

10.23 

11.36 

10.75 

12.39 

1.48 

13.32 

11.93 

11.82 

11.45 

13.09 

1.14 

13.48 

11.49 

11.67 

10.87 

12.90 

1.21 

12.52 

10.76 

11.01 

10.87 

12.92 

1.35 

11.44 

9.57 

10.45 

9.70 

11.92 

1.95 

11.17 

9.35 

9.86 

9.37 

10.82 

1.20 

• 

9.42 

10.09 

9.34 

10.88 

0.87 

9.56 

7.69 

• 

8.00 

9.44 

2.92 

• 

• 

• 

• 

• 

1.16 

12.34 

10.51 

11.25 

10.51 

12.63 

1.16 

10.28 

8.57 

9.00 

9.10 

10.17 


# 

49 

50 

51 

52 

53 

54 

55 

56 

57 

58 

59 

60 

61 

62 

63 

64 

65 

66 

67 

68 

69 

70 

71 

72 

73 

74 

75 

76 

77 

78 

79 

80 

81 

82 

83 

84 

85 

86 

87 

88 

89 

90 

91 

92 

93 

94 

95 

96 


414 


ALVW 

ALPI 

ALPII 

ALPIII 

ALPIV 

ALPV 

0.78 

10.62 

10.32 

• 

10.38 

10.96 

0.50 

11.82 

9.93 

• 

10.13 

11.63 

• 

9.86 

8.48 

8.83 

• 

10.33 

0.63 

10.05 

9.10 


8.72 

9.67 

• 

8.85 

8.90 


9.80 

9.11 

0.59 

9.00 

7.93 


8.06 

9.11 

0.40 

8.45 

7.28 


7.53 

8.08 

• 

7.72 

7.12 


7.09 

7.87 

1.93 

• 

13.93 

14.63 

14.16 

17.37 

1.76 

16.35 

14.13 

13.83 

14.33 

16.49 

• 

22.78 

18.89 

19.12 

19.18 

22.44 

1.81 

15.44 

12.75 

13.22 

12.35 

15.15 

0.91 

13.38 

11.96 

12.01 

11.43 

13.51 

1.53 

17.80 

15.82 

16.38 

15.41 

17.97 

1.75 

13.13 

10.21 

10.12 

9.86 

12.83 

1.52 

14.24 

11.30 

11.79 

11.30 

13.81 

3.93 

29.42 

23.95 

24.75 

23.87 

29.76 

1.27 

15.11 

12.06 

12.72 

11.97 

15.03 

1.85 

17.48 

13.55 

14.24 

13.67 

17.67 

2.74 

20.28 

15.58 

16.71 

15.34 

19.79 

• 

13.75 

11.45 

12.00 

11.17 

• 

2.05 

15.49 

12.58 

14.55 

13.41 

15.97 

1.29 

13.22 

11.32 

12.25 

11.69 

13.30 

• 

24.81 

18.80 

17.28 

18.78 

22.45 

1.21 

15.10 

12.49 

13.43 

13.01 

15.15 

1.02 

14.52 

13.43 

13.58 

13.67 

14.42 

2.13 

16.91 

13.69 

14.10 

13.71 

16.89 

3.23 

32.68 

24.37 

24.43 

26.26 

32.88 

• 

• 

9.90 

10.56 

10.23 

• 

1.94 

12.61 

9.98 

11.08 

10.56 

12.48 

1.14 

11.72 

9.42 

10.12 

9.72 

11.50 

0.53 

10.71 

9.85 

• 

10.08 

10.97 

1.20 

12.21 

10.42 

« 

10.36 

12.03 

• 

10.23 

9.01 

9.43 

8.99 

9.98 

1.10 

12.00 

10.35 

• 

10.75 

12.48 

• 

• 

• 

15.65 

14.52 

16.45 

• 

14.40 

11.40 

11.32 

11.73 

• 

1.62 

13.03 

10.83 

11.11 

11.17 

12.96 

1.10 

17.60 

15.03 

14.68 

15.18 

18.35 

3.26 

31.08 

25.15 

25.46 

25.19 

31.08 

1.38 

10.89 

9.16 

9.38 

9.55 

11.03 

1.07 

12.68 

10.21 

10.47 

10.61 

13.01 

0.96 

11.65 

9.47 

9.62 

9.96 

11.68 

0.76 

10.97 

9.94 

• 

10.10 

10.92 

1.23 

11.53 

9.24 

9.90 

9.75 

11.73 

1.29 

11.25 

8.87 

• 

9.29 

11.09 

0.88 

11.89 

10.02 

10.92 

10.51 

11.92 


# 

97 

98 

99 

100 

101 

102 

103 

104 

105 

106 

107 

108 

109 

110 

111 

112 

113 

114 

115 

116 

117 

118 

119 

120 

121 

122 

123 

124 

125 

126 

127 

128 

129 

130 

131 

132 

133 

134 

135 

136 

137 

138 

139 

140 

141 

142 

143 

144 


415 


ALVW 

ALPI 

ALPII 

ALPIII 

ALPIV 

ALPV 

0.80 

• 

• 

• 

• 

• 

2.70 

30.93 

24.56 

24.77 

25.46 

31.87 

2.43 

23.41 

18.80 

18.88 

19.39 

23.81 

2.81 

27.71 

22.07 

21.94 

22.07 

27.63 

1.82 

• 

• 

• 

• 

• 

1.87 

25.67 

20.81 

21.98 

21.14 

25.71 

1.81 

12.63 

10.16 

10.50 

10.45 

12.87 

• 

10.99 

9.93 

• 

10.26 

• 

1.14 

11.58 

9.56 

9.41 

9.39 

11.62 

0.86 

11.62 

9.72 

« 

9.86 

11.73 

1.27 

11.60 

9.88 

10.47 

10.12 

11.76 

1.11 

10.26 

9.38 

10.26 

10.23 

10.96 

0.62 

10.88 

9.30 

9.60 

9.81 

10.99 

1.48 

12.90 

10.32 

10.71 

10.16 

12.64 

0.88 

11.65 

9.49 

• 

10.21 

12.02 

0.77 

9.37 

7.72 

8.36 

7.59 

9.35 

0.62 

9.58 

8.62 

• 

8.68 

10.08 

0.91 

• 

• 

• 

• 

• 

0.69 

9.34 

8.00 

• 

7.91 

9.39 

1.23 

11.91 

8.99 

9.72 

9.85 

12.56 

0.53 

• 

9.09 

• 

8.14 

• 

• 

8.42 

9.32 

• 

• 

2.51 

• 

26.09 

• 

21.64 

« 

21.04 

22.83 

• 

26.98 

1.94 

19.60 

15.96 

16.38 

16.53 

20.47 

1.21 

11.93 

9.34 

9.27 

9.86 

12.06 

0.67 

10.71 

9.56 

9.25 

9.49 

10.75 

0.57 

10.10 

7.96 

7.97 

8.02 

10.21 

0.93 

9.84 

8.35 

8.96 

8.22 

9.80 

4.41 

32.53 

26.87 

26.41 

26.56 

32.38 

1.19 

15.50 

13.50 

14.17 

13.69 

15.45 

2.84 

29.83 

24.40 

24.71 

24.59 

31.15 

4.80 

41.99 

32.51 

31.82 

33.89 

42.91 

2.46 

24.62 

• 

22.12 

21.15 

25.14 

1.91 

27.07 

23.02 

24.40 

23.21 

28.09 

2.28 

22.07 

19.10 

19.91 

19.12 

22.03 

• 

• 

18.00 

17.62 

17.22 

• 

2.37 

22.18 

19.73 

21.46 

19.58 

22.72 

2.36 

24.15 

21.03 

22.64 

22.22 

23.99 

0.41 

9.65 

8.58 

• 

8.40 

9.33 

1.87 

27.13 

22.40 

22.42 

22.35 

26.78 

1.68 

24.30 

21.74 

22.68 

21.64 

24.89 

2.43 

21.71 

18.59 

19.27 

19.82 

22.91 

0.74 

• 

• 

• 

• 

• 

2.07 

25.24 

21.14 

22.97 

22.52 

25.84 

2.89 

• 

• 

• 

• 

• 

0.76 

10.65 

9.80 

• 

9.82 

10.46 


# 

145 

146 

147 

148 

149 

150 

151 

152 

153 

154 

155 

156 

157 

158 

159 

160 

161 

162 

163 

164 

165 

166 

167 

168 

169 

170 

171 

172 

173 

174 

175 

176 

177 

178 

179 

180 

181 

182 

183 

184 

185 

186 

187 

188 

189 

190 

191 

192 


416 


ALVW 

ALPI 

ALPII 

0.99 

12.06 

10.56 

1.52 

• 

• 

3.08 

28.62 

24.43 

1.82 

26.36 

21.46 

2.07 

43.29 

33.94 

2.48 

39.78 

32.96 

1.80 

37.51 

29.81 

3.35 

46.38 

37.61 

1.86 

30.53 

26.76 

2.55 

39.57 

32.42 

2.73 

44.81 

40.94 

1.20 

30.05 

26.66 

2.22 

35.59 

28.96 

2.79 

41.38 

33.74 

2.04 

31.53 

27.58 

3.29 

44.74 

36.44 

2.40 

38.77 

33.55 

1.73 

34.83 

29.51 

2.28 

31.60 

26.41 

1.99 

33.47 

29.50 

2.87 

44.93 

37.41 

2.50 

47.88 

39.03 

1.93 

36.56 

30.34 

2.28 

33.99 

26.40 

1.70 

35.14 

28.13 

2.19 

33.30 

• 

2.07 

32.06 

27.33 

2.12 

41.20 

34.34 

2.70 

38.73 

34.48 

1.84 

34.47 

29.34 

1.91 

30.59 

25.56 

2.10 

40.34 

34.38 

2.39 

28.70 

24.58 

2.55 

36.48 

30.12 

2.03 

38.18 

33.94 

• 

39.24 

33.56 

1.94 

33.78 

27.89 

1.86 

36.52 

31.01 

1.94 

38.62 

32.83 

1.97 

34.61 

28.96 

2.04 

43.99 

36.81 

1.91 

32.93 

28.71 

2.58 

39.75 

34.03 

2.64 

45.11 

37.90 

2.27 

39.12 

33.32 

2.42 

38.21 

31.18 

2.01 

34.37 

30.60 

1.89 

34.75 

28.01 


ALPIII ALPIV ALPV 

11.58 12.76 

• • • 

23.79 24.05 28.21 

22.09 • 26.73 

35.28 41.41 

34.09 40.06 

30.94 37.85 

36.26 45.22 

26.68 30.13 

32.55 39.16 

38.11 44.29 

• 26.06 30.07 

29.27 36.22 

34.17 41.35 

28.27 32.52 

. 37,11 44.48 

• 34.48 39.61 

• 29.79 35.28 

27.21 31.96 

• 30.46 35.25 

37.56 45.01 

39.58 47.78 

29.92 36.15 

27.99 32.88 

30.81 33.23 

28.01 31.87 

27.57 31.91 

34.91 41.11 

35.71 40.50 

27.96 33.96 

26.25 30.12 

33.33 40.17 

• 24.35 29.36 

30.42 35.85 

34.02 38.99 

33.56 39.72 

27.80 33.66 

31.22 36.84 

34.45 39.65 

29.19 33.41 

38.08 44.36 

27.26 31.90 

• 34.68 40.59 

38.75 45.28 

33.51 38.77 

30.97 37.36 

30.60 35.52 

• 29.07 34.65 


# 

193 

194 

195 

196 

197 

198 

199 

200 

201 

202 

203 

204 

205 

206 

207 

208 

209 

210 

211 

212 

213 

214 

215 

216 

217 

218 

219 

220 

221 

222 

223 

224 

225 

226 

227 

228 

229 

230 

231 

232 

233 

234 

235 

236 

237 

238 

239 

240 


417 


ALWV 

ALPI 

ALPII 

ALPIII 

ALPIV 

ALPV 

2.78 

38.56 

33.85 

• 

34.73 

38.34 

1.73 

26.97 

24.20 

• 

24.30 

26.12 

2.27 

36.26 

30.90 

• 

31.34 

36.55 

2.60 

32.33 

27.22 

• 

28.16 

32.70 

2.26 

38.16 

31.45 

« 

31.29 

38.01 

2.28 

34.82 

31.67 

• 

30.17 

34.13 

2.41 

34.27 

29.98 


30.30 

35.01 

2.46 

38.11 

32.20 

• 

32.11 

37.87 

1.73 

36.42 

30.75 

• 

30.53 

36.16 

2.33 

38.32 

33.80 

• 

34.56 

38.93 

1.49 

27.27 

23.86 

• 

24.61 

27.56 

2.89 

45.22 

39.77 


39.24 

45.23 

1.85 

25.95 

23.69 

• 

23.74 

26.84 

2.78 

50.29 

43.08 

• 

44.17 

51.48 

2.07 

35.34 

29.01 

• 

29.57 

34.95 

2.23 

34.11 

29.65 


29.86 

34.26 

1.71 

23.04 

• 

• 

21.39 

22.87 

1.41 

27.31 

23.08 

• 

22.89 

28.42 

2.21 

36.45 

31.29 

• 

30.57 

36.89 

1.98 

33.78 

28.31 

• 

28.57 

34.33 

2.09 

39.33 

32.29 

• 

32.82 

38.60 

2.25 

39.23 

32.32 

• 

32.31 

39.20 

1.92 

36.58 

30.72 

• 

31.62 

36.55 

1.90 

42.85 

32.74 

• 

33.78 

41.99 

2.18 

37.56 

32.99 

• 

33.66 

37.29 

2.76 

39.49 

34.35 

• 

35.72 

40.04 

1.82 

40.52 

33.95 

• 

34.05 

39.70 

1.59 

32.69 

28.30 


29.24 

32.39 

1.94 

33.08 

28.94 

• 

29.35 

33.38 

2.63 

41.83 

36.40 


36.76 

42.45 

2.09 

37.50 

30.60 


31.88 

38.55 

3.16 

46.29 

40.30 


44.86 

46.64 

2.50 

37.76 

31.73 


34.22 

38.22 

2.34 

25.83 

21.66 

• 

21.95 

25.78 

2.33 

• 

• 



• 

3.00 

43.91 

36.60 


38.73 

44.90 

• 

36.62 

• 

• 

37.57 

45.68 

2.55 

36.96 

29.57 


30.18 

36.66 

2.20 

46.52 

37.98 


37.40 

46.32 

1.72 

33.95 

28.38 

• 

29.09 

34.84 

4.25 

43.33 

34.83 

39.20 

35.24 

43.40 

4.48 

45.19 

38.59 

40.34 

36.80 

45.41 

4.61 

47.15 

37.51 

39.67 

37.68 

46.74 

4.85 

40.44 

34.34 

38.12 

33.54 

42.01 

3.73 

41.50 

33.74 

35.54 

32.62 

40.13 

4.16 

35.92 

26.74 

28.14 

27.05 

36.43 

3.36 

28.63 

24.86 

25.40 

25.06 

29.24 

4.64 

36.01 

29.18 

31.62 

29.59 

35.86 


# 

241 

242 

243 

244 

245 

246 

247 

248 

249 

250 

251 

252 

253 

254 

255 

256 

257 

258 

259 

260 

261 

262 

263 

264 

265 

266 

267 

268 

269 

270 

271 

272 

273 

274 

275 

276 

277 

278 

279 

280 

281 

282 

283 

284 

285 

286 

287 

288 


418 


ALVW 

3.63 

3.40 

2.98 

3.02 

2.78 

2.74 

2.99 

3.03 

3.16 

2.62 

2.74 
• 

2.56 

2.57 

2.41 

2.81 

1.81 

2.01 

2.34 

2.38 

1.75 

2.34 

4.90 

3.71 

2.90 

4.93 

6.86 

6.95 

7.14 

5.63 

5.74 

4.03 

4.71 

3.69 

4.73 

2.65 

3.68 

3.09 

2.89 
3.67 

3.37 

4.06 

4.26 

2.58 

2.77 

3.97 

3.38 
3.11 


ALPI 

30.22 

26.55 

23.26 

22.14 

21.60 

20.00 

20.77 

20.96 

19.08 
18.40 

18.73 

18.37 

18.49 

13.70 

18.44 

16.64 
16.85 

16.67 

16.16 

14.88 

14.78 

11.22 

33.62 
36.10 

26.80 

49.65 

49.33 

55.72 

50.27 

42.63 

43.55 

37.68 

33.96 

30.38 

32.60 

21.74 
32.05 

30.50 

24.56 

34.08 
27.47 
33.94 

33.28 

31.46 
23.76 

31.30 

27.01 

24.46 


ALPII 

22.07 

20.29 

18.50 

18.49 

17.57 

16.89 

16.86 

16.35 

14.92 
15.32 

15.31 

14.93 
14.42 

11.68 

15.49 

14.19 

13.91 

12.77 

12.77 

12.70 

12.31 

9.60 

27.74 

30.07 

21.20 

42.89 
40.52 

45.19 

40.39 

39.01 
39.82 

29.80 

28.36 

26.32 

26.55 

17.91 

28.08 

25.74 

18.78 

27.49 

22.28 
26.35 
26.10 

27.70 

20.14 

25.88 

21.18 

21.25 


ALPIII 

25.06 

22.02 

20.40 

19.90 

18.38 

18.12 

17.46 

17.56 

16.28 

15.62 
17.42 

15.34 

15.49 

10.97 

15.18 

14.16 

14.69 

13.89 

12.63 

12.96 

12.90 
10.04 

28.96 

31.56 

23.47 

44.78 

42.18 

45.92 

44.64 

40.59 

41.49 

32.49 

30.72 

27.59 

29.18 

18.56 

30.95 
27.41 
20.12 

28.30 

23.95 

27.89 

26.29 

28.03 
21.83 

26.21 
24.24 

22.94 


ALPIV 

22.98 

21.18 

19.36 

18.88 

18.50 

17.39 

17.38 

16.91 

15.37 

15.39 

15.72 

15.03 

14.70 

11.86 

15.40 

14.35 
14.28 

13.61 
13.11 

12.80 

12.14 

9.58 

28.65 

31.66 

22.63 

42.78 

39.18 

45.19 

43.30 

39.09 

40.25 

29.10 

28.18 

26.63 
27.82 

18.01 

29.20 

25.61 

18.81 

26.50 

22.38 

26.34 

26.45 

26.60 

20.10 

26.33 

22.31 

21.17 


ALPV 

28.93 

27.17 

23.58 

22.03 

21.97 

20.67 

20.77 

21.10 

19.32 

19.15 

18.32 

18.92 

18.26 

14.27 

18.46 

17.09 

16.62 

17.34 

16.07 

15.10 

14.51 

11.04 

34.82 

36.68 

27.52 

50.29 

49.75 

54.73 

50.86 

48.33 
43.71 

38.04 

33.37 

31.89 

33.04 

21.07 

32.66 

30.93 

24.35 

33.07 

28.01 

34.21 

33.04 
31.44 

24.35 

31.64 

27.38 

24.64 


# 

289 

290 

291 

292 

293 

294 

295 

296 

297 

298 

299 

300 

301 

302 

303 

304 

305 

306 

307 

308 

309 

310 

311 

312 

313 

314 

315 

316 

317 

318 

319 

320 

321 

322 

323 

324 

325 

326 

327 

328 

329 

330 

331 

332 

333 

334 

335 

336 


419 


ALVW 

2.99 

3.08 

3.02 

3.57 

3.07 

3.24 

3.89 

3.15 
2.39 

3.19 

2.74 

2.70 

2.65 
2.69 

2.89 
2.27 

3.14 

2.74 
2.46 

2.77 
3.53 

3.03 

2.43 

2.93 

2.76 

3.45 

2.74 
2.48 

2.57 
2.79 

2.65 

2.33 

2.31 

2.15 

2.77 

2.46 

2.08 

1.94 
2.00 

1.95 
2.00 
1.84 

2.19 

1.66 

1.82 

4.42 

3.19 

4.08 


ALPI 

23.67 

30.47 

24.87 

22.01 

25.30 

22.29 

24.78 

23.19 

20.60 

21.73 
22.26 

19.05 

20.20 

19.17 

19.93 
17.09 

17.80 

17.34 
17.54 

17.80 

20.17 
18.16 

17.63 

19.78 

19.24 

23.13 

16.35 

16.77 

19.60 

16.98 

16.21 
15.34 

15.80 

18.75 
13.86 

15.42 

13.08 

12.06 

12.19 

12.03 

11.63 

11.01 

11.08 

10.98 

10.73 
50.67 

33.02 

30.01 


ALPII 

20.38 

23.14 

19.30 

18.71 

18.99 

18.15 

21.47 

19.07 

17.20 

17.88 

17.93 

16.72 

16.93 

15.63 

15.64 

13.98 

14.45 

14.34 

15.95 

14.82 

16.70 

14.94 

15.25 
15.27 

16.30 

18.64 

13.14 

13.11 

16.76 

14.66 

13.17 

13.03 

12.60 

15.82 

11.80 
12.49 

11.11 

9.89 

10.35 
9.96 
9.65 

9.07 
9.36 

9.07 

8.98 
39.41 

27.08 

23.67 


ALPIII 

20.49 

24.29 

20.47 

19.13 

19.08 

18.45 

22.70 
20.66 
18.58 

19.42 

19.35 

17.04 

17.03 

15.99 

16.67 

14.26 

15.29 

15.02 

15.65 

14.96 

17.24 
16.10 
15.51 

16.09 

18.26 

19.18 

14.01 

14.27 
16.85 

14.60 

13.35 

13.73 

12.65 

15.94 

12.18 

12.96 

11.35 

10.14 
10.75 

10.31 

10.01 

9.09 

9.76 

9.30 
8.91 

41.67 
30.44 

24.77 


ALPIV 

19.33 

23.25 

19.30 

19.20 

18.81 

17.99 

21.42 

19.21 

17.25 

17.92 

17.97 

16.99 

17.57 

15.58 

16.31 

13.89 

14.73 

14.31 

16.15 

14.64 

16.78 

14.81 

14.94 

16.07 

17.02 

18.26 
13.37 

13.35 

16.18 

14.93 

14.13 

13.01 

13.22 

15.67 

12.04 

12.65 

11.09 
10.52 

10.47 

10.45 

9.98 
9.36 

9.74 
9.28 

9.03 

40.67 
27.62 

24.01 


ALPV 

23.08 

29.31 
24.63 

21.43 

25.05 

22.70 

25.21 

23.16 

20.93 

22.13 

23.09 

19.97 

20.98 
19.56 

20.15 

16.90 

18.01 

17.12 

17.81 

17.78 

20.10 
18.62 

17.90 

19.89 

20.12 

22.22 
16.39 
16.92 

19.74 
17.41 

16.10 

15.36 

16.14 
18.80 
13.88 

15.29 

13.10 
12.48 
12.38 

12.47 

13.71 

11.15 

11.34 

11.32 

11.05 

51.11 

33.98 

29.44 


# 

337 

338 

339 

340 

341 

342 

343 

344 

345 

346 

347 

348 

349 

350 

351 

352 

353 

354 

355 

356 

357 

358 

359 

360 

361 

362 

363 

364 

365 

366 

367 

368 

369 

370 

371 

372 


420 


ALWV 

4.07 

3.73 

3.47 

3.69 

2.81 

3.65 
3.34 

2.65 
3.27 

2.50 

2.58 

2.39 

1.83 
1.72 

2.02 

1.60 
1.49 

2.95 
2.42 

2.88 

• 

4.33 

3.51 

• 

4.97 

3.41 

2.24 


4.27 

2.42 


ALPI 

27.74 

25.16 

25.08 

23.70 
24.19 

21.74 

21.73 

21.02 

19.21 

18.63 

16.76 

13.88 

13.43 

13.44 

12.44 

11.17 

8.55 

23.39 


37.56 


26.86 

27.77 

41.37 

37.87 

17.17 

15.22 

14.83 

13.69 

50.88 

33.00 


ALPII 

22.63 

20.84 

20.44 

19.69 
19.21 
20.79 

17.93 

17.25 
16.62 

15.39 

13.78 

11.71 

11.09 

11.69 

10.93 
9.45 

7.07 


29.22 


29.60 

23.63 

24.25 

32.00 

27.31 

12.76 

11.18 
11.51 

10.93 

34.77 

28.46 


ALPIII 

24.13 

21.71 
20.41 
21.83 

20.32 

19.54 

18.33 

18.65 

17.29 

16.60 

14.71 
12.17 
11.57 

11.55 

10.78 
9.61 
7.64 


30.32 


24.35 

• 

30.46 

26.44 

11.98 

11.64 

11.58 

11.01 

33.74 

29.15 


ALPIV 

22.29 

21.19 

20.75 

19.99 

19.34 

21.19 

18.47 

16.98 

16.64 

15.77 

13.67 

12.08 

11.79 

12.04 

10.93 

9.66 

7.23 

20.15 


28.66 


23.55 

31.22 

28.07 

12.31 

11.95 

11.95 

11.29 

36.00 

29.54 


ALPV 

27.42 

26.23 

25.59 

24.03 
23.97 

22.09 
22.18 

21.04 

19.59 

18.84 

16.60 
13.86 

13.74 
14.06 

12.88 

11.29 
8.63 

23.76 


36.74 


37.94 

26.73 

27.85 

41.93 
37.82 

16.94 

14.89 

15.07 

13.65 

51.26 

34.78 


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BIOGRAPHICAL SKETCH 


Craig W. Oyen was born and raised in Williston, ND, where he attended 
public schools and graduated from Williston Senior High School. He may have 
been subliminally directed toward his interest in geology by his father, who gave 
him plenty of “field experience” in the subject. His father allowed him to pick 
rocks from the wheat fields on the family farm (near Zahl, ND) each spring after 
nature’s annual frost-heaving of glacial till boulders to the surface of the fields. 
Another influence may have been a result of activity near his hometown. The oil 
industry was active in the Williston Basin in the late 1970s and early 1980s, and 
Craig occasionally worked as a “swamper,” delivering drilling muds to oil drilling 
rigs in Montana and North Dakota. 

Craig attended North Dakota State University in Fargo, ND, graduating 
with a major in geology and a minor in soil science. It was here that his interest 
in geology and paleontology was strongly influenced by the faculty of the 
Geology Department (particularly Drs. Alan Ashworth and Donald Schwert). 

After his graduation from NDSU, he began graduate studies at the University of 
Tennessee at Knoxville, and became interested in the fossil echinoids of Florida 
while working with Dr. Michael McKinney (a former UF geology graduate student 
of Dr. Douglas Jones). Craig decided to transfer to the University of Florida 


436 


437 


for easier access to the fossil collection in the Florida Museum of Natural History 
and his field locations in the state of Florida. During his graduate work at the 
University of Florida he was employed as a graduate teaching assistant in the 
Department of Geological Sciences, and as a graduate research assistant in 
Florida Museum of Natural History (Invertebrate Paleontology Division) and the 
Department of Geological Sciences. As part of his teaching assistantship, he 
taught laboratory and/or lecture sections of Exploring the Geological Sciences, 
Physical Geology, Environmental Geology, Sedimentary Geology, Invertebrate 
Paleontology, Field Methods in Geology, and Clay Mineralogy. He also gave 
occasional lectures in several other courses. His research appointments 
involved fieldwork and fossil collection, preparation, and cataloguing for the 
FLMNH, and thin-section petrography and analysis of rocks to be used as 
concrete aggregates by the Florida Department of Transportation. 

During the latter stages of his dissertation work, Craig was employed as a 
temporary Assistant Professor in the Department of Geology and Geography at 
Georgia Southern University, in Statesboro, Georgia. He currently is employed 
as an Assistant Professor in the Department of Geography and Earth Science at 
Shippensburg University of Pennsylvania, in Shippensburg, Pennsylvania. 

Craig enjoys participating in many athletic activities such as cycling and 
basketball, though he has slowed down somewhat in recent years. During his 
time as a graduate student at the University of Florida he participated in 
intramural sports, usually on teams composed dominantly of geology graduate 
students (the “Psychotic Basement Trolls”). As a result of these “non-contact” 


438 


sports, he also kept the medical staff at Shands Hospital and the university 
infirmary busy repairing sports injuries including problems such as a cornea 
abrasion (basketball); a level-2 AC joint separation of his shoulder (football); 
numerous patella dislocations (basketball, racquetball); a minor ACL tear in knee 
(basketball); a chipped distal femoral condyle (basketball); a fragmented patella 
(basketball); a broken radius (basketball); a relocated tibia tubercle (and an 
associated tibia fracture and bone screws via reconstruction) (basketball); and 
numerous cuts and abrasions (basketball, football, cycling). Fortunately, with 
age comes wisdom, and he currently exercises more cautiously than before. 

Craig is a member of the following professional organizations and honor 
societies: the Paleontological Society, the Geological Society of America, the 
Florida Paleontological Society, the Pennsylvania Academy of Science, Sigma 
Xi, Sigma Gamma Epsilon, and Phi Kappa Phi. He also serves on the board of 
directors of the Marine Science Consortium, Wallops Island, Virginia. 


I certify that I have read this study and that in my opinion it conforms to 
acceptable standards of scholarly presentation and is fully adequate, in scope 
and quality, as a dissertation for the degree of Doctor of Philosophy. 



I certify that I have read this study and that in my opinion it conforms to 
acceptable standards of scholarly presentation and is fully adequate, in scope 
and quality, as a dissertation for the degree of Doctor of Philosophy. 



I certify that I have read this study and that in my opinion it conforms to 
acceptable standards of scholarly presentation and is fully adequate, in scope 
and quality, as a dissertation for the degree of Doctor of Philosophy. 


Bruce J^acFadden 
Professor of Geology 

I certify that I have read this study and that in my opinion it conforms to 
acceptable standards of scholarly presentation and is fully adequate, in scope 
and quality, as a dissertation for the degree of Doctor of Philosophy. 



Professor of Geology 


I certify that I have read this study and that in my opinion it conforms to 
acceptable standards of scholarly presentation and is fully adequate, in scope 
and quality, as a dissertation for the degree of Doctor of Philosophy. 



This dissertation was submitted to the Graduate Faculty of the Department 
of Geological Sciences in the College of Liberal Arts and Sciences and to the 
Graduate School and was accepted as partial fulfillment of the requirements for 
the degree of Doctor of Philosophy. 

May, 2001 


Dean, Graduate School